;^§#^f*|*l**rJr
The Philippine
Journal of Science
Volume 46
SEPTEMBER TO DECEMBER, 1931
WITH 62 PLATES AND 129 TEXT FIGURES
MANILA
BUREAU OF PRINTING
1931
EDITORIAL BOARD
William H. Brown, Ph.D., Editor
R. C. McGregor, A.B., Associate Editor
Lucilb M. Lidstone, Copy Editor
Chemistry
A. P. West, Ph.D.; T. Dar Juan, Phar.D.
A. S. Arguelles, B.S.; F. D. Reyes, B.S.; R. H. Aguilar, Ch.E.
J. C. Espinosa, B.S. in Ch.E.; Manuel Roxas, Ph.D.
Maria Y. Orosa, Ph.C, M.S.
Geology
Victoriano Elicano, B.S.; Leopoldo A. Faustino, E.M., Ph.D.
Experimental Medicine
Otto Schobl, M.D.; H. W. Wade, M.D.
Stanton Youngberg, D.V.M.; Arturo Garcia, M.D.
Daniel de la Paz, M.D.; Cristobal Manalang, M.D.
Clinical Medicine
Ijiborio Gomez, M.D., Ph.D.; F. Calderon, L.M.
Jacobo Fajardo, M.D.; Jose Albert, M.D.; H. Lara, M.D.
Jose Rodriguez, M.D.; Carmelo Reyes, M.D.
Botany
L. M. Guerrero, Phar.D.; A. F. Fischer, C.E., M.F.
J. K. Santos, Ph.D.; P. L. Sherman, Ph.D.; Eduardo Quisumbing, Ph.D.
Joaquin Maranon, Ph.D.; Rafael B. Espino, Ph.D.
H. Atherton Lee, M.S.
Zoology
Heraclio R. Montalban, M.A.; Leopoldo B. Uichanco, Sc.D.
Marcos A. Tubangui, D.V.M.; Manuel D. Sumulong, M.S., D.V.M.
Anthropology
H. 0. Beyer, M.A.; Otto Johns Scheereu, M.A.
E. E. Schneider, B.L.
<~0
CONTENTS
No. 1, September, 1931
[Issued August 21, 1931.]
Page.
Santos, Irene de, Augustus P. West and P. D. Esguerra. Philip-
pine pine-needle oil from Pinus insularis (Endlicher) 1
Two plates.
Alexander, Charles P. New or little-known TipulidaB from the Phil-
ippines (Diptera), X 9
Three plates.
Manalang, C. Origin of the irritating substance in mosquito bite.... 39
One plate.
Manalang, C. Malaria transmission in the Philippines. III. Den-
sity and infective density of Anopheles funestus Giles 47
Gee, N. Gist. Fresh-water sponges of the Philippine Islands 61
Four text figures.
Skvortzow, B. W. Plankton diatoms from Vladivostok Bay 77
Two plates.
Skvortzow, B. W. Mycetozoa from North Manchuria, China 85
Five plates.
Skvortzow, B. W. Pelagic diatoms of Korean Strait of the Sea of
Japan 95
Ten plates.
Rodriguez, Jose, and Fidel C. Plantilla. The histamine test as an
aid in the diagnosis of early leprosy 123
Lopez, A. W. The fly Eutrixopsis javana Townsend (Diptera, Ta-
chinidae), a parasite of the beetle Leucopholis irrorata in Occi-
dental Negros, Philippine Islands 129
Cruz, Aurelio O., and Augustus P. West. Composition of Philip-
pine kapok-seed oil 131
One plate.
Sumulong, Manuel D. The skeleton of the timarau 141
Three plates and four text figures.
No. 2, October, 1931
[Issued September 7, 1931.]
Colet-Vazquez, Ana. Rat-bite fever in the Philippines 159
Three plates.
Sch5bl, Otto. An interpretation of the laws of Brown and Pearce
that govern the course of treponematoses 169
iii
iv Contents
Page.
Schobl, Otto. Coexistent infection with yaws and syphilis 177
Schobl, Otto. The prospects of vaccination and vaccine therapy in
treponematoses 183
Humphrey, C. J. Decay of wood in automobiles in the Tropics 189
Two plates.
Cruz, Aurelio 0., and Augustus P. West. Composition of Philip-
pine peanut oil 199
One plate.
Copeland, Edwin Bingham. New or interesting Oriental ferns 209
Hadden, F. C, and A. W. Lopez. Efforts toward biological control
of the common pink mealybug Trionymus sacchari (Cockerell)
of sugar cane on Negros 221
Monserrat, Carlos. The Kahn test in clinical syphilis 225
Monserrat, Carlos. Comparative serologic study of Vernes, Was-
sermann, and Kahn reactions in experimental treponematoses 241
Manalang, C. Malaria transmission in the Philippines, IV : Meteo-
rological factors 247
Three text figures.
Santos, Jose K. Leaf and seed structure of the Philippine Coriaria 257
Four plates.
Alexander, Charles P. New or little-known Tipulidae from the
Philippines (Diptera), XI 269
Three plates.
No. 3, November, 1931
[Issued October 15, 1931.]
Russell, Paul F. Avian malaria studies, I. Prophylactic plasmo-
chin in inoculated avian malaria 305
Nineteen text figures.
Russell, Paul F. Avian malaria studies, II. Prophylactic plasmo-
chin versus prophylactic quinine in inoculated avian malaria.... 347
Seven text figures.
Manalang, C. Malaria transmission in the Philippines, V. On
the maturation of the ova of Anopheles funestus Giles 363
One text figure.
Manalang, C. Malaria transmission in the Philippines, VI. The
dark-night factor 371
Cole, Howard Irving. Causes of irritation upon injection of io-
dized ethyl esters of Hydnocarpus-group oils 377
Kleine, R. Die Brenthiden der Philippinen-lnseln 383
Sixteen text figures.
Alexander, Charles P. New or little-known Tipulidae from the
Philippines (Diptera), XII 447
Two plates.
Contents v
Page.
Baker, C. F. Second supplement to the list of the lower fungi of
the Philippine Islands. A bibliographic list chronologically
arranged, and with localities and hosts 479
No. 4, December, 1931
[Issued November 12, 1931.]
Tubangui, Marcos A. Worm parasites of the brown rat (Mus nor-
vegicus) in the Philippine Islands, with special reference to
those forms that may be transmitted to human beings 537
Nineteen text figures.
Holt, R. L., and J. H. Kintner. Notes on dengue 593
Two text figures.
Holt, R. L., Wm. D. Fleming, and J. H. Kintner. Resistance of
dengue virus 601
Four text figures.
Oliver, Wade W., Walfrido de Leon and Alfredo Pio de Roda. The
attempted cultivation of Mycobacterium leprae 611
One plate.
Mendiola, N. B. Somatic segregation in double Hibiscus and its in-
heritance 627
Three plates and one text figure.
Russell, Paul F. Daytime resting places of Anopheles mosquitoes
in the Philippines: First report 639
Four plates.
Russell, Paul F. Avian malaria studies, III. The experimental
epidemiology of avian malaria; introductory paper 651
Two plates and three text figures.
Yeager, Clark H. Bored-hole latrine equipment and construction.... 681
Seven plates and forty-six text figures.
King, W. V. The Philippine varieties of Anopheles gigas and Ano-
pheles lindesayi 751
Two plates.
Lopez, A. W. The use of the antennae as a means of determining the
sexes in Leucopholis irrorata adults (Coleoptera, Scarabaeidse 759
One plate.
Index 765
The Philippine
Journal op Science
Vol. 46 SEPTEMBER, 1931 No. 1
PHILIPPINE PINE-NEEDLE OIL FROM PINUS
INSULARIS (ENDLICHER)
By Irene de Santos and Augustus P. West
Of the Bureau of Science, Manila
and
P. D. Esguerra
Of the Bureau of Forestry, Manila
two plates
Benguet pine (Pinus insularis Endlicher) forms extensive for-
ests in the mountain regions of northern Luzon. Some months
ago we investigated samples of resin tapped from the Benguet
pine and our results showed that a turpentine x of good quality
and a high-grade rosin 2 are obtained from it. Recently we in-
vestigated the oil obtained from the leaves (needles) of Ben-
guet pine and found that the yield of oil is very small. The
oil appears to consist largely of alpha- and beta-pinene and to
contain only a small percentage of esters calculated as bornyl
acetate.
Leaf oils are obtained from the leaves of numerous kinds of
trees. Oil distilled from the leaves of pine trees is known as
pine-needle oil and a number of these oils obtained from dif-
1 Santos, I. de, A. P. West, and J. Fontanoza, Philip. Journ. Sci. 45
(1931) 233.
2 Santos, I. de, A. P. West, and J. Fontanoza, loc. cit.
259737 1
2 The Philippine Journal of Science 1931
ferent species of pine have been investigated. In general, these
oils consist essentially of a mixture of terpenes.
Pine-needle oil has a fragrant odor and is useful in com-
pounding perfumes. It has also been employed as a repellent
for certain insects 3 and as a larvicide for mosquitoes.4
EXPERIMENTAL PROCEDURE
Through the kindness of Mr. Sixto Laraya, of the Philippine
Bureau of Forestry, our laboratory has been supplied during
recent months with occasional shipments of pine-needles and
twigs. These were gathered from pine trees growing in and
near Baguio, a summer resort situated at an elevation of about
1,500 meters in Mountain Province, Luzon.
The pine-needles and twigs were placed in a large appara-
tus and steam distilled. The pine-needle oil thus obtained was
separated from the aqueous distillate and dehydrated with
calcium chloride. The yield of oil distilled from leaves and
twigs was only 0.043 per cent and subsequent experiments
showed that most of the yield was obtained from the leaves and
not from the leafless twigs. The oil was slightly greenish yel-
low in color and had a strong aromatic odor. The constants of
the oil were determined and the data are recorded in Table 1.
Table 1. — Constants of pine-needle oil.
Specific gravity (*——} 0.8582
(QQOX
A "d P desrees +20.53
Refractive index (u j?J 1.4700
Acid value 1.38
Saponification value 7.67
Ester value (7.67 — 1.38) 6.29
Esters as bornyl acetate, per cent 1.75
Benguet pine-needle oil was found to be soluble in 10 parts
of alcohol (90 per cent). When distilled (fractionated) we ob-
tained the results recorded in Table 2. The first three frac-
tions were practically colorless while the residue had a red
color.
3 Bishop, F. C, R. C. Roark, D. C. Parman, and E. W. Laake, Journ.
Econ. Entomol. 18 (1925) 776.
* Barnes, M. E., Am. Journ. Hyg. 5 (1925) 309.
46,1
Be Santos et ah: Pine-needle Oil
Table 2. — Distillation of Benguet pine-needle oil (Amount of oil distilled,
76 cubic centimeters.)
Fraction.
Refractive
index,
XT30oC
N-D~-
Specific
gravity,
,300c
d"ioc-.
Optical
rotation,
A30oC
A"D-
(100 mm.
tube).
No.
Temperature.
Amount obtained.
1
2
3
4
Below 155
155 to 160
Grams.
3.7
34.4
26.7
10.8
Per cent.
4.9
45.3
35.1
14.2
Degrees.
1.4645
1.4677
0.8476
0.8493
0.9420
+28.7
+ 18.8
160 to 164. _ _
Residue . . .
In Table 3 are given the boiling points of a few terpene com-
pounds that commonly occur in pine-needle oils.
Table 3. — Boiling points of a few common terpenes.
Terpene.
Alpha-pinene
Beta-pinene
Dipentene
Limonene
Borneol
Boiling
point, ° C.
156-157
164-166
170-172
172.6-178.2
208-213
A comparison of the data given in Tables 2 and 3 indicates
that fraction 2 of Benguet pine-needle oil probably contains
alpha-pinene and fraction 3 beta-pinene. A sample of fraction
2 was cooled in ice and treated with dry hydrochloric acid gas.
There separated out a heavy thick oil that had a strong odor
of pinene hydrochloride (artificial camphor). The presence of
other substances seemed to prevent the hydrochloride from
crystallizing.
A portion of fraction 3 was oxidized with alkaline permanga-
nate. The reaction product was steam distilled and the res-
idue filtered to eliminate manganese oxide. When the filtrate
was evaporated somewhat and cooled, white crystals of sodium
nopinate separated out. A portion of these crystals was de-
composed with dilute sulphuric acid and extracted with ben-
zene. Needles melting at 125.5° to 127° C. were thus obtained,
indicating that fraction No. 3 contains beta-pinene.
Due to the very small yield of oil from Benguet pine leaves
we did not have sufficient material to make a very thorough
investigation of the composition of Benguet pine-needle oil.
The Philippine Journal of Science
1931
In Table 4 are given the constants of pine-needle oils from
various species of pine. The figures for specific gravity, refrac-
tive index, and optical rotation are not exactly comparable since
they were determined at somewhat different temperatures.
As shown by the data (Table 4) pine-needle oils from dif-
ferent species of pine vary considerably in composition. With
the exception of Pinus insularis and P. sylvestris, these oils list-
ed below give a negative rotation. In general, the yield of
pine-needle oils is very small. Only one oil (Pinus pumilis) gave
a yield of more than 0.5 per cent while the other oils gave
considerably less.
Table 4. — Constants of pine-needle oils from different species of pine.
Species.
Pinus insularis tt
Abies sibirieah
Do
Pinus longifolia b
Pinus pumilio b
Do
Pinus sabiniana b_ . .
Do
Abies magnified b
Pinus contorta b
Pinus ponderosa h. . _ .
Do
Pinus lambertiana b_
Do
Pinus halepensis °__ .
Picea vulgaris d
Do
Pinus sylvestris e
Pinus exceha {
Yield
Per cent.
0.043
0.250
0.750
0.078
0.102
0.040
0.126
0.045
0.120
0.260
0.310
Specific
gravity.
0.8582
0.9000
0.9280
0,8740
0.8630
0.8760
0.8510
0.8570
0.8665
0.869C
0.8718
0.8849
0.8676
0.8738
0.8960
0.8800
0.8880
0.8661
0.8672
Optical
rotation.
+20.53
—30.00
—43.00
— 6.15
— 5.00
—10.00
—20.00
—39.00
— 16.7C
—17.84
—15.73
—19.59
—11.07
—16.50
—49.44
—21.70
—37.00
4-13.20
—13.76
Refractive
index.
1.4700
1.4700
1.4730
1.4740
1.4800
1.4670
1.4671
1.4861
1.4831
1.4789
1.4838
1.4777
1.4794
1.4940
1.4729
1.4727
Acid
value.
1.38
1.00
4.00
1.03
1.47
2.05
0.75
0.90
0.67
2.36
0.68
2.38
0.28
1.00
Esters as
bornyl
acetate.
Per cent.
1.75
29.00
43.00
5.00
3.00
10.00
2.37
3.32
3.47
2.11
1.36
2.83
0.74
2.07
6.58
2.90
3.43
3.75
a Pinus insularis from the Philippines.
b Parry, E. J., Chemistry Essential Oils and Perfumes 1 (1918) 51.
c Rutovskii, B. N., Perfum. Essen. Oils. Rec. 19 (1928) 391.
d Allen's Commercial Organic Analysis 4 (1925) 112.
e Rao, B. S., and J. L. Simonsen, Journ. Chem. Soc. 127 (1925) 2494.
f Rutovskii, B., I. Vinogradova, and V. Koslov., Arbeiten Chem. Pharm. Inst. Moskaus,
Lief 11 (1925) 93.
The authors wish to thank Mr. Arthur F. Fischer, director,
Philippine Bureau of Forestry, and Mr. Luis J. Reyes, chief,
division of forest products, Bureau of Forestry, for their co-
operation and assistance in this work.
46, i De Santos et ah : Pine-needle Oil 5
The authors also wish to thank Mr. Sixto Laraya, of the
Philippine Bureau of Forestry, for his kindness in procuring
samples of Benguet pine needles for this investigation.
SUMMARY
We have investigated the pine-needle oil obtained from Ben-
guet pine (Pinus insularis Endl.).
Benguet pine-needle oil has a positive optical rotation and in
this respect is unlike most pine-needle oils which have a negative
rotation.
Compared to other pine-needle oils, the yield from Benguet
pine leaves is very small (0.043 per cent).
Benguet pine-needle oil appears to consist largely of alpha-
and beta-pinene and to contain only a small percentage of es-
ters calculated as bornyl acetate. It is soluble in ten parts of
90 per cent alcohol.
ILLUSTRATIONS
Plates 1 and 2. Philippine pine trees in Baguio.
De Santos and West: Pine-needle Oil.]
[Philip, Journ. Scl» 46, No, 1.
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NEW OR LITTLE-KNOWN TIPULIDJE FROM THE
PHILIPPINES (DIPTERA), X1
By Chakles P. Alexander
Of Amherst, Massachusetts
THREE PLATES
The important series of Philippine crane flies discussed at
this time were collected in Luzon by Messrs. McGregor, Duyag,
and Rivera, and in Mindanao by Mr. Charles F. Clagg. I wish
to thank the above-mentioned gentlemen for their continued
interest in making known the tremendously rich tipulid fauna
of the Philippines. All types are preserved in my collection.
TIPULIN^E
SCAMBONEURA NIGROTERGATA sp. nov. Plate 2, fig. 23.
General coloration obscure yellow; antennae (male) elongate,
the scapal segments yellow?; mesonotal praescutum with three
narrow, ill-delimited, reddish brown lines; postnotal medioter-
gite and pleura yellow, unmarked; wings subhyaline; anterior
arculus bowed ; abdominal tergites with a continuous black dor-
somedian stripe from base to apex ; sternites light yellow ; male
hypopygium with the tergite uniformly blackened; appendage
of ninth sternite small, bilobed.
Male. — Length, about 13 millimeters; wing, 11.3; antenna,
about 7.
Frontal prolongation of head obscure yellow; nasus black;
palpi light brown, the outer segment passing into black. An-
tennae (male) elongate, as shown by the measurements; scape
obscure yellow; flagellum black, the segments elongate, their
longest verticils about one-fourth to one-fifth the segment. Head
obscure orange, with a brown median line on vertex ; additional
narrower and less-defined dark lines on vertex, delimiting the
posterior vertex.
Pronotum obscure yellow. Mesonotal praescutum obscure yel-
low, with three narrow reddish brown stripes that are ill-de-
1 Contribution from the entomological laboratory, Massachusetts Agri-
cultural College.
9
10 The Philippine Journal of Science 1931
limited; scutum yellowish testaceous, the cephalic half of the
lobes blackened; scutellum testaceous; postnotal mediotergite
yellow, unmarked. Pleura yellow. Haiteres brownish black,
the knobs black. Legs with the coxae and trochanters yellow;
femora brownish black, their bases broadly yellow; tibiae and
tarsi black. Wings subhyaline, iridescent, the stigmal region
dark brown; veins black. Venation: Anterior cord strongly
bowed ; m-cu nearly half its length beyond the fork of M.
Abdomen with the tergites pale, with a continuous dull black
median stripe the entire length, more extensive and somewhat
paler on outer segments; a narrower continuous lateral black
line ; sternites clear light yellow ; hypopygium yellow, the tergite
entirely black. Male hypopygium (Plate 2, fig. 23) with the
tergite, 9t, bearing two conspicuous earlike lobes, separated by a
V-shaped median notch that further bears a tiny median tongue-
like projection; mesal margin of lobes with delicate setulae at
apex, these replaced by coarse black setae that merge gradually
into short black spines on the face of the lobes. Outer dististyle,
od, obliquely broadest beyond base, the outline irregular, the
outer edge most protuberant just beyond base, the inner margin
more strongly rounded at near midlength. Appendage of ninth
sternite, 9s, small, conspicuously bilobed, the entire surface se-
tiferous.
Mindanao, Davao district, Calian, La Lun Mountains, alti-
tude 5,500 feet, July 5, 1930 (Clagg) ; holotype, male.
This species and the next are very different from the other
known species of Scamboneura, although closely allied to one
another. The nearest ally in Luzon would seem to be S. vitti-
vertex Alexander.
SCAMBONEURA CALIANENSIS sp. hoy. Plate 2, fiffs. 24 and 25.
Male. — Length, about 15 millimeters; wing, 14.2; antenna,
about 6.
Generally similar to S. nigrotergata sp. nov. ; in the general
coloration, differing as follows :
Size larger, but the antennae (male) proportionately and ac-
tually shorter, as shown by the measurements, the flagellar seg-
ments being conspicuously shorter. Scutal lobes with the
markings reddish brown and occupying the whole lobe. Pleura
yellow, vaguely marked with more reddish yellow on the anepis-
ternum and ventral sternopleurite. Abdomen with the dorso-
median black stripe not quite continuous, being narrowly
46fi Alexander: Philippine Tipulidse, X 11
interrupted at the caudal margins of the segments. Male hy-
popygium with the tergite (Plate 2, fig. 24, 9t) entirely
blackened and shaped generally as in nigrotergata but the details
quite different. Lateral ears conspicuous, with abundant long
coarse setae but no replacement spines on disk; median projec-
tion large and conspicuous. Outer dististyle (Plate 2, fig. 25)
long and conspicuous, the apex produced into a slender point.
Mindanao, Davao district, Calian, La Lun Mountains, altitude
5,500 feet, July 4, 1930 (Clagg) ; holotype, male.
LIMONIIN^E
LIMONIINI
LIMONIA (LIMONIA) CANDIDELLA sp. nor. Plate 1, fig. 1; Plate 2, fig. 26.
General coloration yellow, the praescutum with a brown me-
dian stripe; antennae black, the flagellar segments cordate, with
glabrous apical necks; legs black, all tarsi with intermediate
portions white; wings with a strong blackish suffusion; male
hypopygium with the basistyles elongate, the ventromesal lobe
small, at extreme base.
Male. — Length, about 6 millimeters; wing, 6.8.
Rostrum and palpi black. Antennae black throughout; inter-
mediate flagellar segments cordate, with glabrous apical necks
that are about one-third the length of the segment; outer seg-
ments more elongate; terminal segment long, about one-half
longer than the penultimate ; verticils shorter than the segments.
Head black, the front silvery ; anterior vertex relatively wide.
Pronotum dark medially, obscure yellow on sides. Mesonotal
praescutum clear yellow, with a narrow median brown stripe,
the usual lateral stripes ill-delimited, brownish yellow, the hu-
meral region brightest ; scutum yellow, the centers of the lobes
vaguely darker; scutellum brown, obscurely brightened poste-
riorly; postnotum brownish yellow. Pleura light yellow, the
dorsal sclerites a little more darkened. Halteres black. Legs
with the coxae and trochanters pale yellow, the fore coxae a
trifle more darkened; femora brownish black, the bases re-
strictedly brightened; tibiae dark brown; tarsi dark brown, the
intermediate portion of all tarsi snowy white, this involving the
distal third or more of basitarsi, the entire second segment and
all but the tip of the third segment; the amount of white greatest
on the hind legs where the distal two-thirds of the basitarsus
is included ; claws small, with a single basal tooth. Wings (Plate
1, fig. 1) with a strong black suffusion, the small oval stigma
12 The Philippine Journal of Science 1931
darker; extreme wing tip vaguely darkened; veins brownish
black. Costal fringe of moderate length. Venation: Sc long,
Scx ending beyond the fork of Rs, Se2 a short distance from its
tip ; free tip of Sc2 and R2 in alignment ; Rs less than twice the
basal section of R4+5; cell 1st M2 closed, shorter than any of
the veins beyond it; m-cu just beyond the fork of M; vein 2d
A long, converging strongly toward 1st A.
Abdominal tergites black ; basal sternites obscure yellow ; outer
sternites and hypopygium darker. Male hypopygium (Plate 2,
fig. 26) much as in L. multinodulosa in the median extension of
the tergite, elongate basistyles, b, with the ventromesal lobe small
and situated at the extreme base, and the general conformation
of the dististyles and gonapophyses. The dorsal dististyle, dd,
is a more strongly curved hook.
Luzon, Laguna Province, above Ube, foot of Mount Banahao,
altitude about 700 meters, on mossy cliff near river in cool for-
est, February 9, 1930 (McGregor) ; holotype, male.
Limonia (Limonia) candidella is most closely allied to L. (L.)
multinodulosa Alexander (Luzon), differing in the much shorter
antennae of the male and the white intermediate tarsal segments
of all the legs. The increased length of the antennae in multi-
nodulosa is produced by the longer glabrous apical necks of the
segments.
LIMONIA (LIMONIA) LATIFLAVA sp. nov. Plate 1, fig. 2.
General coloration brownish yellow, the posterior sclerites of
the mesonotum blackened; antennae black; pleura yellow with
a black longitudinal stripe; halteres black; legs black, the tarsi
and broad tibial tips yellowish white; wings with a blackish
tinge, the basal cells streaked with whitish; Sc long, Sc2 at tip
of Scx.
Female. — Length, about 5 millimeters; wing, 5.
Rostrum black, relatively long and conspicuous, about one-
half as long as the remainder of head; palpi black. Antennae
black throughout ; flagellar segments oval. Head black, sparse-
ly pruinose; anterior vertex narrow, lighter gray.
Pronotum black. Mesonotal praescutum obscure brownish yel-
low, paler laterally, more brownish imedially ; posterior sclerites
of mesonotum more uniformly brownish black. Pleura with a
conspicuous longitudinal black stripe extending from the pro-
notum to the abdomen, the dorsopleural region obscure yellow;
ventral pleural region clear light yellow. Halteres black. Legs
46, i Alexander: Philippine Tipulidse, X 13
with the coxae and trochanters light yellow ; femora black, their
bases restrictedly pale ; tibiae black, the tips paling to yellowish
white, this subequal in amount on all legs and including about
the distal fourth or fifth ; tarsi similarly yellowish white. Wings
(Plate 1, fig. 2) with a strong blackish suffusion, the oval stigma
darker ; conspicuous whitish streaks in the proximal ends of cells
R, M, Cu, and both anals ; veins brownish black. Venation : Sc
long, Scx ending shortly before the fork of Rs, Sc2 at its tip ; Rs
long, arcuated; free tip of Sc2 and R2 in transverse alignment;
m-cu just beyond the fork of M; vein 2d A converging toward
1st A at base.
Abdominal tergites brownish black, the sternites yellow. Ovi-
positor with the tergal valves relatively small, strongly upcurved ;
sternal valves straight, their bases blackened.
Luzon, Laguna Province, above Ube, February, 1930 (McGre-
gor) ; holotype, female.
Limonia (Limonia) latiflava is very different from other re-
gional species of the subgenus, the most distinctive characters
being the very extensive pale apices of all the legs, the color-
ation involving not only the entire tarsi but also the tips of the
tibiae.
LIMONIA (LIMONIA) FLAVOHUMERALIS sp. nov. Plate 1, figf. 3; Plate 2, fiff. 27.
General coloration black, the humeral and lateral regions of
the prsescutum broadly and conspicuously light yellow; pleura
with a broad black longitudinal stripe; wings dark gray, the
margins still darker; Sc long, Sc2 at tip of Scx; male hypopy-
gium with the ventral dististyle small, the rostral prolongation
long, without spines.
Male. — Length, about 3.5 millimeters; wing, 4.2.
Rostrum and palpi black. Antennae black; flagellar segments
oval, passing into more elongate-oval ; terminal segment elongate,
about one-half longer than the penultimate, the distal end point-
ed; verticils short. Head large, especially the eyes; dorsum
dark gray, the anterior vertex reduced to a capillary strip.
Pronotum black, the posterior notum yellow. Mesonotal praes-
cutum light yellow, including the very broad humeral and lat-
eral portions; a triangular brownish black median shield on
posterior half, this sending a scarcely apparent vitta cephalad
to the margin; scutal lobes blackened, the median area testa-
ceous, the lateral margins yellow; scutellum brownish black;
postnotal mediotergite testaceous brown. Pleura with the dorsal
14 The Philippine Journal of Science 1931
portion occupied by a broad black longitudinal stripe that ex-
tends from the pronotum to the abdomen, encircling the root
of the halteres; sternopleurite and meral region pale yellow;
dorsopleural region adjoining the wing root obscure yellow.
Halteres infuscated. Legs with the fore coxae darkened, the
other coxae and all trochanters yellow; femora dark brown, the
bases narrowly obscure yellow; remainder of legs brownish
black, the tarsi very insensibly paler; claws nearly simple.
Wings (Plate 1, fig. 3) with the disk dark gray, the margins
more infuscated ; stigma subcircular, darker brown ; conspicuous
dusky seams along vein Cu in cell M, along Rs and the cord;
veins brownish black. Costal fringe short; macrotrichia of
veins long and conspicuous. Venation: Sc long, Scx ending
about opposite two-thirds the length of Rs, Sc2 at its tip; cell
1st M2 closed, relatively short ; m-cu close to fork of M ; cell 2d
A narrow, the veins gently converging near origin.
Abdomen brownish black; hypopygium dark. Male hypopy-
gium (Plate 2, fig. 27) with the tergite, dt, transverse, the
caudal margin convexly rounded, with a deep and narrow me-
dian incision. Basistyle, b, relatively large, especially the large,
obtuse, ventromesal lobe. Dorsal dististyle a short, stout, flat-
tened blade, the apex suddenly narrowed to an acute point.
Ventral dististyle small, oval, much smaller than the basistyle,
the body of the style with long coarse setae ; rostral prolongation
long and slender, without rostral spines. Gonapophyses, g,
with the mesal-apical angle a stout lobe.
Luzon, Laguna Province, above Ube, at foot of Mount Bana-
hao, altitude about 700 meters, in cool forest, February 9, 1930
(McGregor) ; holotype male.
Limonia (Limonia) flavohumeralis is most similar in general
coloration to L. (L.) retrusa Alexander (Luzon), differing very
notably in all details of coloration and structure of the male
hypopygium.
LIMONIA (LIMONIA) CANIS sp. nov. Plate 1, fig. 4; Plate 2, fig. 28.
Allied to L. cynotis; general coloration dark brown; wings
with a strong blackish tinge, without stigmal darkening; free tip
of Sc2 far before R2; male hypopygium with the ventromesal
lobe of basistyle long and slender ; dististyle single, shaped more
or less like a dog's ear, the mesal face on apical half with spi-
nous setae.
46,1 Alexander: Philippine Tipulidse, X 15
Male. — Length, about 4 millimeters ; wing, 4.6.
Rostrum and palpi dark brown. Antennae black throughout;
basal flagellar segments subglobular, passing to oval outwardly ;
terminal segment scarcely longer than the penultimate ; segments
densely clothed with microscopic black setulae and a few stout
verticils of moderate length. Head dull black.
Mesonotum chiefly brownish black, the pleura paler, more
obscure testaceous. Halteres infuscated. Legs with the coxae
and trochanters testaceous yellow; remainder of legs dark
brown. Wings (Plate 1, fig. 4) with a strong blackish suffusion,
without a stigmal darkening; veins darker brown. Venation : Sc
long, Scj. ending at about two-thirds to three-fourths the length
of the nearly straight Us, Sc2 at its tip; free tip of Sc2 far
before level of R2; m-cu close to fork of M; anal veins nearly
parallel to very weakly convergent at origin.
Abdomen brownish black. Male hypopygium (Plate 2, fig.
28) generally as in L. cynotis in the conformation of the styli,
differing conspicuously in details. Tergite, 9t, large, narrowed
outwardly, the caudal margin with a broad U-shaped emargina-
tion. Basistyle, 6, with the ventromesal lobe very long and
slender, only a little shorter than the dististyle, narrowed out-
wardly. Dististyle, d, single, shaped more or less like a dog's
ear, the spinous setae on mesal face restricted to distal half.
Gonapophyses, g, pale, the mesal-apical lobe slender, the tip pro-
duced slightly laterad into a point.
Luzon, Laguna Province, above Ube, at foot of Mount Ba-
nahao, altitude about 700 meters, near river in cool forest,
February 9, 1930 (McGregor) ; holotype, male.
Limonia (Limonia) canis is allied to L. (L.) cynotis Alexander
(Mindanao), differing most evidently in the structure of the
male hypopygium, especially the long ventromesal lobe of the
basistyle and the vestiture of the dististyle.
LIMONIA (RHIPIDIA) MORIONELLA (Edwards).
Rhipidia (Rhipidia) morionella Edwards, Journ. Fed. Malay States
Mus. 14 (1928) 70.
Luzon, Mountain Province, Benguet, Mount Santo Tomas, al-
titude over 5,000 feet, March 21 to 24, 1930 (Rivera) ; Tayabas
Province, Candelaria, June 25, 1930 (McGregor and Rivera) .
These agree exactly with the types from the Federated Malay
States except that the second tarsal segment is darkened.
16 The Philippine Journal of Science 1931
LIMONIA (RHIPIDIA) LUTEIPLEURALIS sp. nov. Plate lt fisr. 5.
Belongs to the rostrifera group ; closely allied to L. morionella;
general coloration black, the thoracic pleura yellow, only the
ventral sternopleurite darkened; wings unmarked except for
stigma; terminal tarsal segments whitish.
Male. — Length, about 3.5 to 3.8 millimeters; wing, 4 to 4.4.
Female. — Length, about 4 millimeters; wing, 3.8 to 4.
Closely allied to L. morionella; differing especially in the
yellowish thoracic pleura.
Rostrum longer than the remainder of head, black. Antennae
black, the apices of the axial portions of the segments paler ; an-
tennae of male long-bipectinate ; of female, simple. Head black.
Mesonotum brownish black, the pleura obscure yellow, only
the ventral sternopleurite darkened. Halteres dusky, the base
of stem restrictedly pale. Legs with the coxae and trochanters
yellow; remainder of legs black, the femoral bases restrictedly
brightened; subterminal tarsal segments restrictedly whitish,
more extensive and clearer white on posterior legs where from
one-fourth to one-third of the tarsus is this color. Wings
(Plate 1, fig. 5) whitish hyaline, unmarked except for the con-
spicuous short-oval brown stigma; veins brownish black. Ve-
nation: SCi ending about opposite one-third the length of Rs,
Sc2 far from its tip, Scx alone being one-half longer than Rs;
cell M2 open by the atrophy of m; cell 2d A wide.
Abdominal tergites dark brown, the sternites more yellow.
Male hypopygium dark brown. Ovipositor with the genital
shield blackened, the valves paling to horn-color.
Luzon, Mountain Province, Benguet, Mount Santo Tomas, al-
titude over 5,000 feet, March 21 to 25, 1930 (Rivera) ; holotype,
male; allotype, female; paratypes, numerous males and females.
Although closely allied to L. (R.) morionella (Edwards), I
must consider the present fly to be distinct by reason of the
yellow thoracic pleura. The amount of white on the tarsi is
more restricted and obscured in the present species. It should
be observed that following the inclusion of Rhipidia as a sub-
genus of Limonia (Limnobia), morionella Edwards (1928) be-
comes preoccupied by morionella Schiner (1868) and should be
renamed. The members of the rostrifera group do not seem to
be strictly consubgeneric with Rhipidia but rather to represent
a distinct off -shoot of the genus.
46,1 Alexander: Philippine Tipulidse, X 17
LIMONIA (GERANOMYIA) PHCENOSOMA sp. nov. Plate 1, fig. 6; Plate 2, fig. 29.
General coloration reddish ; head blackish gray with a silvery
median vitta ; postnotal mediotergite dark brown ; knobs of hal-
teres blackened ; wings with a faint brown tinge, sparsely marked
with small brown clouds that are distributed in the costal field ;
male hypopygium with the cephalic margin of the rostral pro-
longation of ventral dististyle with sclerotized bracing areas;
rostral spines very elongate ; gonapophyses with apices of mesal-
apical lobes bifid.
Male. — Length, excluding rostrum, about 5.5 millimeters;
wing, 5.6; rostrum, about 2.2 to 2.3.
Female. — Length, excluding rostrum, about 5.5 to 7 milli-
meters ; wing, 5.5 to 6.3 ; rostrum, about 2.4 to 2.8.
Rostrum and palpi black, the former of moderate length only,
slightly longer in the female. Antennae black throughout; fla-
gellar segments oval to subcylindrical, the verticils short and
inconspicuous. Front and anterior vertex silvery ; remainder of
head blackish gray, with a silvery median vitta to the occiput;
anterior vertex narrow.
Mesonotum shiny reddish yellow, the disk of the prsescutum
and the scutal lobes darker, more chestnut-red, the lateral por-
tions more yellowish; scutellum obscure yellow, darker basally;
postnotal mediotergite conspicuously dark brown, the lateral por-
tions yellow. Pleura reddish yellow. Halteres pale, the knobs
inf uscated. Legs with the coxae and trochanters yellow ; femora
obscure yellow, more brownish on distal half; tibiae and tarsi
brownish yellow, the terminal segments darkened; claws with
a powerful basal tooth, with an additional microscopic denticle
more proximad. Wings (Plate 1, fig. 6) with a faint brown
tinge, sparsely patterned with brown, including the stigma and
small spots at origin of Rs, fork of Sc, along anterior cord, and
as a marginal seam in the radial field; narrow and less con-
spicuous seams to the supernumerary crossvein in cell Sc, along
posterior cord, and on outer end of cell 1st M2 ; veins brownish
black. Costal fringe short. Venation: Sc long, Scx ending at
near four-fifths the length of Rs, Sc2 at its tip ; a supernumerary
crossvein in cell Sc at about two-thirds the length of vein R ; Rs
weakly angulated at origin ; cell 1st M2 closed ; m-cu at or before
the fork of M ; cell 2d A narrow, the anal veins at base generally
parallel.
259737 2
18 The Philippine Journal of Science iwi
Abdominal tergites brownish black, the basal segments a little
brightened laterally at the incisures; sternites yellow; outer
segments of abdomen paler in both sexes. Male hypopygium
(Plate 2, fig. 29) with the tergite, 9t, transverse, the caudal
margin convexly rounded, divided by a small median notch into
two halves that are provided with abundant setae. Basistyle,
6, relatively small, the ventromesal lobe large, conspicuously seti-
ferous. Dorsal dististyle a very strongly curved chitinized
sickle, the acute tip blackened. Ventral dististyle, vd, fleshy,
oblique, the conspicuous rostral prolongation protected along
its cephalic margin by sclerotized areas ; two very long, curved,
rostral spines, arising from a common basal tubercle, placed
near apex of the prolongation. Gonapophyses, g, with the mesal-
apical lobes conspicuously bifid at apex.
Luzon, Laguna Province, above Ube, February 6 to 12, 1930
(McGregor and Rivera) ; holotype, male ; allotype, female ; para-
type, female; Tayabas Province, Candelaria, June 25, 1930
(McGregor and Rivera) ; paratype, female.
Limonia (Geranomyia) phcenosoma is readily told by the pe-
culiar structure of the male hypopygium.
LIMONIA (GERANOMYIA) LONGIFIMBRIATA sp. nov. Plate 1, fig. 7; Plate 2, fig. 30.
General coloration yellow; mesonotal praescutum with three
gray stripes that are separated by two narrow blackish lines;
halteres dusky ; wings with a faint brownish tinge, very sparsely
patterned with brown ; costal fringe (male) very long and con-
spicuous; cell M2 open by the atrophy of m; male hypopygium
with the ventral dististyle large and fleshy ; spines of the rostral
prolongation from long basal tubercles that are widely separated ;
mesal-apical lobe of gonapophyses very long and slender.
Male. — Length, excluding rostrum, about 6 siillimeters ;
wing, 6 to 6.2 ; rostrum, about 2.5.
Female. — Length, excluding rostrum, about 5 millimeters;
wing, 6.2; rostrum, about 2.
Rostrum and palpi black. Antennae black throughout; fla-
gellar segments cylindrical with short inconspicuous verticils.
Head blackish gray; a narrow silvery line from the front to the
occiput.
Pronotum blackish gray. Mesonotal praescutum with three
gray stripes, the interspaces dull black, the humeral and lateral
regions obscure yellow ; scutum obscure yellow, the lobes exten-
sively blackish gray; scutellum testaceous; postnotal medioter-
46fi Alexander: Philippine Tipvlidse, X 19
gite dark brown, especially on the posterior half. Pleura ob-
scure yellow, the pleurotergite a trifle darkened. Halteres dus-
ky. Legs with the coxae and trochanters pale greenish yellow;
remainder of legs dark brown, the femoral bases restrictedly
brightened; basal tarsal segments paling to brownish yellow.
Wings (Plate 1, fig. 7) with a faint brown tinge; stigma oval,
dark brown; a vague gray clouding along cord; wing apex in
radial field narrowly bordered by brown; veins dark brown.
Costal fringe (male) very long and conspicuous. Venation: Sc
long, SCi ending about opposite or beyond midlength of Rs,
Sc2 at its tip ; a supernumerary crossvein in cell Sc ; cell M2 open
by the atrophy of m; m-cu at or close to the fork of M.
Abdominal tergites dark brown, the sternites greenish yellow.
Male hypopygium (Plate 2, fig. 30) with the caudal margin of
tergite gently emarginate, with two low lobes. Basistyle, 6,
relatively small, the ventromesal lobe moderately large. Ven-
tral dististyle, vd, a very large fleshy lobe, the rostral prolonga-
tion large, complex in structure, the two spines arising from
widely separated pale tubercles, the inner spine shorter. Dor-
sal dististyle a strongly curved pale sickle, the tips slightly up-
curved. Gonapophyses, g9 with the mesal-apical lobe very long
and slender, gently curved.
Luzon, Laguna Province, Mount Maquiling, May 23 to 30,
1930 (Duyag) ; holotype, male; paratype, male; above Ube, al-
titude 400 meters, January 27, 1930 (McGregor), paratype,
male; Pampanga Province, Mount Arayat, October, 1929 (Ri-
vera) ; allotype, female.
Limonia (Geranomyia) longifimbriata is very distinct from
regional species in the unusually long costal fringe in the male
and the open cell M2.
LIMONIA (GERANOMYIA) PARAMANCA sp. nov. Plate 1, fi*. 8.
Belongs to the argentifera gropp; allied to L. manca in the
open cell M2 ; wings with a strong dusky tinge, the stigma and
a broad marginal seam in cell R2 darker brown.
Female.— Length, excluding rostrum, about 4.8 millimeters;
wing, 4.5; rostrum, about 2.5.
Rostrum long, black ; palpi black. Antennae black throughout,
the verticils short. Head gray, the front and anterior vertex
silvery ; central portion of posterior vertex extensively blackened.
Mesonotum polished black, the prsescutum with a silvery area
on sides behind pseudosutural foveae, with a smaller similar
20 The Philippine Journal of Science 1931
sublateral area at suture; scutellum and postnotum more prui-
nose. Pleura heavily silvery pruinose, the sternal region paler.
Halteres yellow. Legs with the coxse and trochanters yellow;
femora obscure yellow, slightly darker beyond base ; remainder
of legs brown. Wings (Plate 1, fig. 8) with a strong dusky
tinge, the stigma and a broad marginal seam in cell E2 darker
brown ; veins brownish black. Costal fringe relatively long and
conspicuous for the female sex. Venation: Scx ending beyond
midlength of Rs; an unusually wide supernumerary crossvein
in cell Sc at near two-thirds the length of vein R ; cell M2 open
by the atrophy of m ; m-cu about one-half its length beyond the
fork of M, the distal section of CUi very short ; cell 2d A narrow.
Abdominal tergites black, the sternites paler, more brownish;
genital segment brownish yellow. Ovipositor with the tergal
valves very slender, gently upcurved, reddish horn color.
Luzon, Tayabas Province, Candelaria, June 25, 1930 (McGre-
gor and Rivera) ; holotype, female.
Limonia (Geranomyia) paramanca is readily distinguished
from the other members of the argentifera group by the dark-
ened wings, in conjunction with the small size and open cell
M2. The other members of the group, with the exception of
manca Alexander (North Queensland) have cell 1st M2 closed
(argentifera de Meijere, nigronotata Rrunetti, nigronitida
Alexander, and pleuropalloris Alexander). As I have in-
dicated in another paper, a study of the type specimen of sor-
billans (Wiedemann) shows that it, too, belongs to this group
and is very probably identical with argentifera. The type is
a female, in relatively poor condition, and the synonymy cannot
be readily affirmed.
LIMONIA (PSEUDOGLOCHINA) ANGUSTAPICAUS sp. nov. Plate 1, fig. 9; Plate 2,
figr. 31.
General coloration dark brown, the pronotum and broad pleu-
ral region yellowi; fore femora white, the tips narrowly black-
ened; posterior femora dark brown; remainder of legs snowy
white, all tibiae with a single narrow black ring at midlength;
wings whitish, the large stigma and narrow apex blackened;
abdominal sternites distinctly bicolored ; male hypopygium with
a single stout rostral spine.
Male. — Length, about 5 millimeters ; wing, 5.5.
Female. — Length, about 5 millimeters; wing, 5.
Rostrum pale yellow; palpi black. Antennae black through-
out, relatively elongate, the long-oval segments with short apical
pedicels. Head yellow, more dusky on the orbits.
46,1 Alexander: Philippine Tipulidse, X 21
Pronotum light yellow. Mesonotum dark brown, with a more
or less distinct paler median line from the posterior portion of
the praescutum to the postnotal mediotergite where it becomes
more pruinose. Pleura chiefly occupied by a broad yellow lon-
gitudinal stripe, more pruinose on its ventral portion; dorsal
pleurotergite and ventral sternopleurite dark brown. Halteres
pale, the knobs infuscated. Legs with the coxae and trochanters
brownish yellow ; fore femora white, the tips narrowly blackened ;
posterior femora dark brown, the tips narrowly blackened; all
tibiae snowy white with a single narrow black ring at midlength ;
tarsi snowy white. Wings (Plate 1, fig. 9) whitish, the apex in
outer radial cells darkened; stigma large, dark brown; veins
black, the prearcular veins R whitish. Venation: Scx ending
just beyond the fork of the short oblique Rs; cell 2d M2 deep;
m-cu at fork of M; cell 2d A small, as in L. unicinctipes.
Abdominal tergites dark brown, the intermediate segments
with a paler brown subterminal area; subterminal segments
blackened; sternites bicolored, the bases broadly black, the tips
about equally yellowish white; ventral dististyle pale yellow.
Male hypopygium (Plate 2, fig. 31) with the ventral dististyle,
vd, large and fleshy, the rostral prolongation small, with a single
short powerful spine. Gonapophyses broad-based, the mesal-
apical angle small.
Luzon, Laguna Province, Mount Maquiling (Duyag) ; holotype,
male; allotype, female, January 28, 1930; paratype, male, May
23 to 30, 1930 (Duyag).
Limonia (Pseudoglochina) angustapicalis is most closely allied
to L. (P). unicinctipes Alexander, differing most conspicuously
in the large stigmal area, distinctly darkened apex of the wings,
and the dimidiate abdominal sternites.
LIMONIA (ALEXANDRIARIA) SOLLICITA sp. nov. Plate 1, &g. 10.
General coloration gray; antennae black throughout; knobs of
halteres dark brown; legs yellow, the terminal three tarsal seg-
ments dark brown ; wings gray, sparsely patterned with brown ;
Sc short, Scx very long ; a marginal spur of vein M3 persisting.
Female. — Length, about 5 millimeters; wing, 5.
Rostrum black, the labial palpi brown, the maxillary palpi
black. Antennae black throughout; flagellar segments oval,
more elongate outwardly, the terminal segment a little longer
than the penultimate. Head gray ; anterior vertex narrow.
Pronotum and mesonotum brown, the three praescutal stripes
darker brown but almost concealed by yellowish pollen; scutal
22 The Philippine Journal of Science 1931
lobes brownish black, the median area paler ; scutellum brownish
gray; postnotum dark gray. Pleura gray. Halteres short, ob-
scure yellow, the knobs dark brown. Legs with the coxae
brownish yellow, the fore coxae somewhat darker; trochanters
yellow ; remainder of legs yellow, the three terminal tarsal seg-
ments infuscated; third and fourth tarsal segments on flexor
surface with rows of evenly spaced pale spines on the entire
length of the segment ; claws small, with a single well-developed
tooth. Wings (Plate 1, fig. 10) gray, sparsely patterned with
brown; stigma oval, brown; restricted grayish brown clouds
at Sc2, origin of Es, and along cord ; veins dark brown. Vena-
tion : Sc short, Scx ending opposite the origin of Rs, very long,
Sc2 being at near midlength of R; a marginal spur of M3 back
from wing edge; m-cu close to fork of M; cell 2d A wide.
Abdominal tergites dark brown, the sternites brownish yellow.
Ovipositor with the tergal valves very slender, the sternal valves
correspondingly stout and deep.
Luzon, Laguna Province, Ube, December, 1929 (Rivera) ;
holotype, female.
Lim<kiia (Alexandricuria) sollicita is very different from the
other regional species of the subgenus. The wing pattern is
almost as in L. (Dicranomyia) sordida (Brunetti) and similar
species. It is uncertain as to how constant the presence of the
marginal vein M3 will prove to be.
ORIMARGA (ORIMARGA) RUBRICOLOR sp. nov. Plate 1, fi*. 11.
General coloration red; antennae black throughout; wings
milky gray, the veins pale; macrotrichia of veins relatively
sparse, there being only about four on the distal half of R3.
Male. — Length, about 3.2 millimeters; wing, 3.8.
Female. — Length, about 3.6 millimeters; wing, 3.8.
Rostrum and palpi black. Antennae black throughout; fla-
gellar segments subglobular, passing into oval outwardly. Head
gray.
Thoracic dorsum reddish brown, the pleura clearer red. Hal-
teres pale. Legs with the coxae reddish ; trochanters testaceous ;
remainder of legs pale brown, long and slender. Wings (Plate
1, fig. 11) milky gray, the prearcular and costal regions light
yellow; veins pale. Costal fringe relatively long and conspic-
uous. Macrotrichia of veins relatively sparse, there being four
on distal half of R3, widely separated ; a series of about twenty
to twenty-five the entire length of the distal section of R4+5,
more crowded toward outer end ; additional trichia on outer half
46,1 Alexander: Philippine Tipulidse, X 23
of each of veins M1+2 and M3. Venation: Scx ending about
opposite three-fifths the length of Rs, Sc2 not far from its tip;
R3 a trifle shorter than R2+3 ; basal section of R4+5 about twice
R2+3; m-cu opposite the proximal third of Rs.
Abdomen entirely red in male, the subterminal segments of
female blackened.
Luzon, Tayabas Province, Candelaria, June 25, 1930
{McGregor and Rivera) ; holotype, male ; allotype, female.
"This red fly is found on damp mossy rocks at streamside." —
McGregor.
Orimarga rubricolor is readily told by the conspicuous red
coloration of the body.
HELIUS (RHAMPHOLIMNOBIA) RETICULARIS (Alexander).
Rhampholimnobia reticularis Alexander, Proc. U. S. Nat. Mus. 49
(1915) 169-170.
One male, Pakawan, Ifugao Subprovince, Mountain Province,
Luzon, April 7, 1930 (Rivera). The species and the subgenus
are new to Luzon and the Philippines, having previously been re-
corded only from Java (type locality) and Borneo.
HEXATOMINI
EPIPHRAGMA (POLYPHRAGMA) BAKERI Alexander. Plate 1, fig. 12; Plate 2, fi*. 32.
Epiphragma bakeri Alexander, Philip. Journ. Sci. 21 (1922) 373-374.
A male from Pauai, Mountain Province, Luzon, altitude 8,000
feet, April 11, 1930 (Rivera), is generally similar to the holotype
male except in the more-restricted brown wing pattern. The
venation (Plate 1, fig. 12) has never been shown. The male
hypopygium (Plate 2, fig. 32) is very different from that
of the other Luzon species of the subgenus so far described.
Region of the tergite, 9t, produced medially into a shield-shaped
area, the caudal margin of which is deeply notched. Basistyle,
b, with a small fleshy lobe on mesal face at base. Interbasal
process, i, expanded on basal half, the apex unequally bidentate.
Outer dististyle, od, a small bottle-shaped structure, the apex bent
at a right angle into two subequal teeth. Inner dististyle, id,
larger, flattened. iEdeagus large, with an irregular elevated
crest.
EPIPHRAGMA (POLYPHRAGMA) PARVILOBA sp. nov. Plate 1, fig. 13; Plate 2, fig. 33.
Male. — Length, about 6.5 to 7 millimeters ; wing, 7.5 to 8.
Generally similar to E. (P.) ochrinota Alexander in the general
coloration of the body, differs most conspicuously in the dark
24 The Philippine Journal of Science 1931
antennae, narrow anterior vertex, wing pattern, and details of
the male hypopygium.
Antennal scape black, the fusion segment infuscated, in cases
a little brightened beneath. Head brownish gray, the anterior
vertex very narrow, the eyes unusually large.
Mesonotum fulvous, contrasting markedly with the black pleu-
ra. Femora yellow, the subterminal darkening relatively pale
and ill-defined. Wings (Plate 1, fig. 13) grayish, the costal
region light yellow; a diffuse brown pattern, darker and more
clearly delimited along the costal margin, the markings of the
disk not bordered by yellow, as is the case in E. ochrinota.
Male hypopygium (Plate 2, fig. 33) with the median tergal
lobes, 9t, very small, separated by a broad U-shaped notch.
Basistyles very long and slender. Outer dististyle, od, dilated at
midlength, setif erous, thence narrowed to an acute curved point,
with a small lateral tubercle before apex. Interbasal process,
i, a long simple spine, more slender than in ochrinota. Phallo-
some, p, with the sedeagus set in a deep notch in the quadrate
plate.
Luzon, Laguna Province, above Ube, foot of Mount Banahao,
altitude 400 to 700 meters, February 3 to 9, 1930 (McGregor
and Rivera) ; holotype, male ; paratypes, 3 males. The holotype
was taken at 700 meters, in flight near river in cool forest.
LIMNOPHILA (EPHELIA) IGOROTA sp. nov. Plate 1, fig. 14; Plate 3, fi«r. 34.
Antennal scape black, the flagellum chiefly pale; mesonotal
prsescutum yellow, with abundant dark markings ; knobs of hal-
teres blackened ; femora yellow, the tips more yellowish brown,
with a very narrow black subterminal ring; wings broad in
male, the dark pattern compact; seam on m-cu narrow, discon-
nected with the major area on the anterior cord.
Male. — Length, about 5.5 millimeters; wing, 6.5
Female. — Length, about 7 millimeters; wing, 7.5
Antennae with the scapal segments dark brown, the first seg-
ment pruinose; flagellum with the basal six to eight segments
light yellow. Head yellow, mottled with blackish.
Mesonotal praescutum with the ground color yellow, the usual
stripes much dissected; lateral stripes entire, connected at an-
terior ends with the pseudosutural fovese and confluent laterally
with the broad dark brown lateral margins of the sclerite;
median prsescutal stripe blackened behind the level of the pseu-
dosutural fovese, the anterior portion wider, more grayish yellow,
mottled with darker dots and with a capillary black vitta; in-
46,i Alexander: Philippine Tipulidse, X 25
terspaces behind the pseudosutural f ovese with four or five dots
that are in part confluent. Pleura gray, with numerous con-
spicuous brown spots that scarcely assume the form of a stripe.
Halteres with the knobs black. Legs with the femora yellow,
the tips light yellowish brown, the proximal end of this dark-
ened ring narrowly blackened, as in L. granulata. Wings (Plate
1, fig. 14) of male broader than in female; dark pattern more
restricted to the costal half, especially of the area along the
cord, which forms an almost solid mass that extends back to the
fork of M, the clear area in cell C greatly restricted, not reach-
ing any of the veins of Rs; the very narrow seam along m-cu
is not connected with the mark along the anterior cord, the Y-
shaped figure in granulata thus appearing more V-shaped ; seam
on m-cu not in alignment with the anterior cord, being at or
beyond midlength of cell 1st M2; dark seam on the supernum-
erary crossvein in cell M a little distad of the general level of
the dark areas that form the first crossband; dark spot beyond
the prearcular area very small and inconspicuous.
Male hypopygium with the apical notch of the outer dististyle,
od (Plate 3, fig. 34), broad and shallow, the margin irregular,
the outer apical angle a decurved spine, preceded by a group
of from five to seven smaller appressed spines ; on lateral margin
of style at near midlength with a conspicuous appressed spinous
lobe.
Luzon, Mountain Province, Benguet, La Trinidad, below Ba-
guio, altitude 4,800 feet, in open parklike area, March 26, 1930
(Rivera) ; holotype, male; allotype, female; Mount Santo Tomas,
above Baguio, altitude over 5,000 feet, March 21, 1930 (Rivera) ;
paratype, female ; Pauai, April 21, 1930 (Rivera) ; paratype,
1 male; Laguna Province, above Ube, altitude 1,500 feet, Feb-
ruary 11, 1930 (Rivera) ; paratypes, 2 males.
Limnophila (Ephelia) igorota is closely allied to the Bornean
L. (E.) granulata Edwards, differing especially in the details of
wing pattern and venation, the black knobs of the halteres, and
other details.
PILARIA PHCENOSOMA sp. nov. Plate 1, fig. 15; Plate 3, fig. 35.
General color red ; antennae short in both sexes ; halteres black ;
wings with a strong brown tinge ; vein R3 very short, not exceed-
ing one-third the length of the long R4, cell R3 at margin thus
being very wide; cell Mx lacking.
Male. — Length, about 7.5 to 8.5 millimeters; wing, 7 to 8.5.
Female. — Length, about 10 millimeters; wing, 7.5.
26 The Philippine Journal of Science 1931
Rostrum and palpi black. Antennae short in both sexes;
scapal segments reddish brown; flagellum black; flagellar seg-
ments short and crowded, the outer segments passing into cylin-
drical; all segments with long conspicuous verticils that exceed
the segments. Head fiery orange; vertex broad.
Thoracic dorsum fiery reddish orange, the prsescutum without
distinct stripes except a vague median capillary darkening;
pseudosutural foveae extensive but pale reddish and so incon-
spicuous ; tuberculate pits at cephalic margin of sclerite reddish ;
scutellum brownish testaceous. Pleura reddish, vaguely marked
with darker on the anepisternum and sternopleurite, the poste-
rior sclerites more testaceous. Halteres black, the extreme base
of stem brightened. Legs with the coxae and trochanters ob-
scure yellow; femora obscure brownish yellow, the tips narrowly
blackened; tibise and tarsi black. Wings (Plate 1, fig. 15) with
a strong brownish tinge, the small oval stigma darker brown;
prearcular and costal regions a little brighter, especially before
and beyond the stigma; conspicuous longitudinal hyaline oblit-
erative streaks in cells R, R3, M, 1st M2, M3, and M4 ; veins dark
brown. Venation: Sc relatively long, Scx ending opposite the
fork of Rs, Sc2 some distance from its tip, Scx alone exceeding
R 4 ; R8 very short, not exceeding one-third the length of
the long R4, cell R3 at margin, thus being very wide; cell M±
lacking; m-cu about one-third to one-half its length beyond the
fork of M ; anterior arculus preserved.
Abdomen reddish, the caudal margins of the tergites narrowly
but conspicuously blackened ; hypopygium orange-yellow. Male
hypopygium (Plate 3, fig. 35) with the tergite, 9t, conspicuous,
the median portion of the caudal margin produced into a broad
lobe that is further produced into two submedian glabrous plates,
their tips obtuse, these plates separated by a deep notch. Basi-
styles, b, short and stout. Dististyles, id, od, as figured, the
inner style very broad.
Luzon, Laguna Province, Ube, February 11 to May 9, 1930
(McGregor and Rivera) ; holotype, male; allotype, female; para-
topotypes, 15 of both sexes.
Pilaria phoenosoma is very different from all described mem-
bers of the genus, in some respects more resembling a small
Eriocera. The following notes on the occurrence of this species
are of much interest: "The water supply for Majayjay comes
from a large spring near Ube. The overflow runs off in a small
stream and is used for irrigation. Just below the spring is
46,1 Alexander: Philippine Tipnlidse, X 27
a small bog (area approximately one hectare). Some of this
is open, with growth of ferns, sedges, and small shrubs. A
large part is covered with a bamboo and pandan thicket. Many
of the mountain streams are dry this month (March), but this
spring seems to have the same overflow as in the rainy months.
In ferns and other low vegetation along this stream and in
plants on this boggy area, many large and small tipulids oc-
curred."— McGregor. Associated with the Pilaria in this hab-
itat on March 4, 1930, were the following Tipulidae: Limonia
(GercCnomyia) argentifera (de Meijere), L. (Goniodinetira) ni-
griceps (van der Wulp), L. (Thrypticomyia) apicalis (Wiede-
mann), Conosia irrorata (Wiedemann), Trentepohlia (Trente-
pohlia) trentepohlii (Wiedemann), Gonomyia (Lipophleps) bi-
colorata Alexander, and Erioptera (Erioptera) rubripes Alex-
ander.
PILARIA CARBONIPES sp. nov. Plate 1, fi*. 16.
General coloration of mesonotum polished black, the thoracic
pleura abruptly yellow; antennae (male) elongate; halteres and
legs black; wings with a blackish tinge; R2 shorter than R2+3;
cell Mx present; hypopygium black.
Male. — Length, about 4 to 4.2 millimeters; wing, 4.3 to 5;
antennae, 2.3 to 2.6.
Female.— Length, about 5 millimeters; wing, 4.6 to 5.
Antennse (male) elongate, much exceeding one-half the length
of the body, black throughout; flagellar segments cylindrical to
elongate-fusiform, with dense erect black setae and slightly
longer verticils. In the female the antennse are shorter, about
equal to the combined head and thorax, the setae lacking or in-
conspicuous, the verticils very long and evident. Head polished
black.
Mesonotum polished black, the humeral region of prsescutum
very restrictedly pale. Pleura, including the pleurotergite,
yellow. Halteres blackened. Legs with the coxae and trochan-
ters obscure yellow ; remainder of legs black, only the femoral
bases restrictedly obscure yellow. Wings (Plate 1, fig. 16) with
a strong blackish tinge, the oval stigma slightly darker brown ;
veins brownish black. Venation: Sc3 ending about opposite
four-fifths the length of Rs, Sc2 at its tip; R2+3 present, a little
longer than R2 alone ; R3 long, straight or weakly sinuous ; inner
ends of cells R4, R5, and 1st M2 in oblique alignment, the last
most basad ; cell M± slightly longer than its petiole ; m-cu beyond
midlength of cell 1st M2.
28 The Philippine Journal of Science mi
Abdominal tergites and hypopygium black, the sternites
abruptly light yellow.
Luzon, Laguna Province, above Ube, February 11 to April 14,
1930 {McGregor and Rivera) ; holotype, male; allotype, female;
paratypes, 6 of both sexes ; Mount Maquiling, January 28, 1930
(Duyag) ; paratype, female.
Pilaria carbonipes is somewhat similar to the Japanese P.
melanota Alexander, differing in the more-blackened notum,
the black legs, and strongly infumed wings, with the venational
details quite distinct, notably the position of R2 and the course
of R3.
PILARIA CARBONIPES HOLOMELANIA subsp. nov.
As in the typical form, but the pleura and pleurotergite
polished black. The legs, especially the tarsi, paler, the tarsi
fading to yellowish white.
Mindanao, Davao district, Calian, La Lun Mountains, altitude
5,800 feet, July 3, 1930 (Clagg) ; holotype, male; allotype, fe-
male ; paratypes, 1 male, 1 female.
PILARIA ALBOPOSTICATA sp. nov.
Male. — Length, about 5 to 5.2 millimeters; wing, 6; antennae,
2.8 to 3.
Female. — Length, about 5.5 millimeters; wing, 5.5.
Characters much as in P. carbonipes sp. nov., differing as
follows : Legs black, the femoral bases obscure yellow, especially
the posterior femora; tarsi black, the posterior tarsi conspic-
uously whitish yellow, the two terminal segments darkened.
Luzon, Laguna Province, above Ube, February, 1930 {Rivera) ;
holotype, male; paratypes, 3 males ; Mountain Province, Benguet,
La Trinidad, below Baguio, altitude 4,800 feet, March 26 to 28,
1930 {Rivera) ; allotype, female; paratypes, 3 of both sexes.
The conspicuous pale coloration of the posterior tarsi is dis-
tinctive of the species.
ERIOPTERINI
GONOMYIA (LIPOPHLEPS) MAQUILINGIA sp. nov. Plate 1, fig. 17; Plate 3, figr. 36.
General coloration brownish gray ; rostrum orange ; antennae
black throughout; thoracic pleura indistinctly variegated yel-
lowish testaceous and pale brown; legs brownish black; wings
tinged with brownish gray, the stigma only vaguely darker; Sc
short; male hypopygium with a single small subterminal dis-
46, i Alexander: Philippine Tipulidte X 29
tistyle; phallosomic structure terminating in a median organ
shaped like a tuning fork.
Male. — Length, about 3 millimeters ; wing, 3.
Eostrum orange; palpi black. Antennae black throughout.
Head chiefly gray.
Mesonotum brownish gray, the median region of the scutum
slightly paler; posterior callosities of scutal lobes and the scu-
tellum obscure yellow, the median region of the latter at base
darkened ; postnotal mediotergite pruinose. Pleura vaguely pat-
terned with yellowish testaceous and pale brown, the pale color-
ation including the posterior sclerites; dorsal pleurites and
ventral sternopleurite darkened. Halteres dusky, the base of
stem restrictedly brightened. Legs with the coxae and trochan-
ters testaceous-brown ; remainder of legs brownish black. Wings
(Plate 1, fig. 17) tinged with brownish gray, the stigma slightly
and vaguely darkened; veins brown. Venation: Sc short, Scx
ending a short distance before origin of Rs, this distance greater
than the length of the latter; Rs less than two-thirds the an-
terior branch of the same; cell 1st M2 closed; m-cu before fork
of M.
Abdominal tergites brown, paler laterally, the sternites more
uniformly pale. Male hypopygium (Plate 3, fig. 36) with the
apical lobe of basistyle, b, slender. Dististyle, d, single, pale,
much shorter and more slender than the lobe of the basistyle,
provided with about six setae. Phallosomic structure, p, con-
sisting of a pale fan-shaped plate, its caudal margin with four
low crenulate lobes; a further median extension is shaped like
a tuning fork.
Luzon, Laguna Province, Mount Maquiling, May 23 to 30, 1930
(Duyag) ; holotype, male.
Gonomyia (Lipophleps) maquilingia is generally similar to
G. (L). incompleta Brunetti, differing decisively in the very
different male hypopygium.
GONOMYIA (LIPOPHLEPS) INCOMPLETA Brunetti.
Gonomyia incompleta Brunetti, Fauna British India, Dipt. Nemato-
cera (1912) 471-472.
Gonomyia (Leiponeura) insulensis Alexander, Can. Ent. 45 (1913)
286-287.
Luzon, Laguna Province, Ube, February 6, 1930 (Rivera);
Tayabas Province, Candelaria, June 25, 1930 (McGregor and
30 The Philippine Journal of Science 1931
Rivera). This fly has a very extensive range in eastern Asia,
from British India to Japan.
GONOMYIA (LIPOPHLEPS) PALLIDISIGNATA sp. nov. Plate 1, fig. 18; Plate 3, fig. 37.
General coloration brown to grayish brown; basal segments
of flagellum yellow, the outer segments blackened; pleura with
a whitish longitudinal stripe ; legs with the femora pale brown,
the tips whitish, inclosing a very broad black subterminal ring ;
tibiae pale brown, the tips narrowly pale yellow; wings white,
clouded with pale brown; Rs from one-third to one-half longer
than the petiole of cell R3; male hypopygium with three dis-
tistyles.
Male. — Length, about 2.8 millimeters; wing, 3.
Female. — Length, about 3.5 millimeters; wing, 3.5.
Rostrum and palpi black. Antennae with the scape above and
basal two segments of flagellum yellow, the remainder of the
organ blackened. Head white, the center of the vertex exten-
sively blackened.
Pronotum and anterior lateral pretergites white. Mesonotum
brown, varying from reddish brown to dark grayish brown, the
scutal lobes darker; scutellum obscure white, the median area
darkened at base; postnotum dark. Pleura brown to brownish
black, usually blue-gray pruinose, with a narrow, conspicuous,
longitudinal white stripe extending from and including the fore
coxae, passing beneath the halteres, this stripe sometimes ob-
scured or lost. Halteres yellow, the base of the club darkened.
Legs with the fore coxae white, the mid-coxae dark brown, the
posterior coxae dark brown on basal half, white on distal half;
trochanters whitish; femora beyond base pale brown, with a
very broad and conspicuous black subterminal ring, preceded
and followed by narrow white annuli that are less than one-third
the area of the blackened annulus ; tibiae pale brown, the tips nar-
rowly pale yellow; tarsi brown. Wings (Plate 1, fig. 18) with
the ground color white, this including the prearcular, costal,
and apical portions ; remainder of disk clouded with pale brown,
reducing the ground color to areas in both ends of cells R and M,
a more or less distinct crossband beyond the cord, and the outer
ends of cells Cu and 1st A; restricted darker brown areas at
origin of Rs and tip of Sc, stigma, ends of veins R3 and R4,
and along the cord ; veins brown, pale in the ground areas. Ve-
nation : Scx ending opposite or shortly beyond origin of Rs, the
latter unusually long for this subgenus, being about one-third
46,1 Alexander: Philippine Tipididse, X 31
to one-half longer than the straight petiole of cell R3; R3 short
and transverse, R4 strongly arcuated ; m-cu before the fork of M.
Abdomen brownish black, including the hypopygium; caudal
margins of abdominal segments narrowly and indistinctly paler.
Male hypopygium (Plate 3, fig. 37) with three dististyles, the
outer a long, gently curved, blackened rod; intermediate style
very small, appearing as a pale spine ; innermost style long-oval,
terminating in two long setee.
Luzon, Laguna Province, Ube and above, altitude 400 to 700
meters, February 6 to April 14, 1930 (McGregor and Rivera) ;
holotype, male; allotype, female; numerous paratypes of both
sexes.
Although closely allied to G. (L.) nubeculosa de Meijere, I
must regard the present fly as being distinct, differing especially
in the coloration of the legs and wings and the longer Rs. I
do not have a male of nubecvlosa for comparison. The African
species, G. (L.) liberiensis Alexander, G. (L.) noctabunda Alex-
ander, and G„ (L.) sobrina Alexander, are also allied though
separable on venation and structure of the male hypopygium.
Edwards is entirely correct and justified in referring this group
of flies with cell R3 preserved to Lipophleps rather than to the
typical subgenus where they had been placed by other workers.
GONOMYIA (LIPOPHLEPS) ALBOANNULATA sp. nov. Plate 1, fi». 19; Plate 3, &g. 38.
Closely related to G. diffusa; rostrum and palpi black; basal
segments of antennal flagellum pale; thoracic pleura with a
narrow white line ; halteres with darkened knobs ; femora brown-
ish yellow, with a brown subterminal ring, preceded and fol-
lowed by clear yellow; wings unmarked except for a vague pale
brown stigmal area; anterior branch of Rs gently sinuous; male
hypopygium with three dististyles, the intermediate one spinous
at apex, the inner style split into three acute spines.
Male.— Length, about 2.6 millimeters; wing, 3.3.
Female.— Length, about 3 millimeters; wing, 3.2.
Rostrum relatively elongate, about one-half the remainder of
head, black; palpi black. Antennae with the scape dark brown,
the basal flagellar segments pale, the outer segments passing
into dark brown. Head pale, the center of the vertex restrictedly
darkened.
Anterior pronotum whitish, with a darkened median spot;
anterior lateral pretergites whitish. Mesonotum grayish brown,
the pseudosutural f oveae dark brown ; median region of scutum
32 The Philippine Journal of Science mi
and narrow posterior margin of scutellum obscure testaceous;
postnotal mediotergite brownish gray, the anterior lateral angles
broadly yellow. Pleura brownish black on ventral half, this in-
closing a conspicuous white longitudinal stripe, bordered on
either side by blackish ; dorsopleural region buffy, more blackened
in front. Halteres pale, the knobs brown. Legs with the fore
and hind coxae pale, the mid-coxae dark brown; trochanters
yellow; femora brown to yellowish brown, with a broad brown
subterminal ring, preceded and followed by a narrow clearer
yellow ring; tibiae white, the tips narrowly blackened; tarsi
white, the tips dark brown. Wings (Plate 1, fig. 19) grayish,
unmarked except for a vague pale brown stigmal area; prear-
cular and costal regions more yellowish; veins pale brown. Ve-
nation : SCj. ending opposite the origin of the strongly arcuated
Rs ; anterior branch of Rs gently sinuous.
Abdomen of male dark brown, including the hypopygium;
caudal margins of segments conspicuously ringed with pale;
pleural membrane conspicuously whitened. In female, the
segments are uniformly darkened, as in diffusa. Male hypopy-
gium (Plate 3, fig. 38) with three dististyles, the outermost a
simple blackened blade, gradually narrowed to the obtuse tip;
intermediate style a little shorter, appearing as a straight rod,
the distal third slightly expanded into a spinous head; inner-
most style, id, trifid, all arms acute, the laterals straight and
provided with two or three setae, the central arm curved, gla-
brous.
Luzon, Tayabas Province, Candelaria, along margin of stream,
June 25, 1930 (McGregor and Rivera) ; holotype, male. Min-
danao, Davao district, Calian, Lawa, at trap lantern, April 24,
1930 (Clagg) ; allotype, female; paratype, female.
Gonomyia (Lipophleps) alboannulata is most closely allied to
G. (L.) diffusa (de Meijere), differing especially in the darkened
knobs of the halteres, the details of venation, as the strongly
sinuous anterior branch of Rs, and the pattern of the legs and
wings. I do not know the male sex of diffusa.
GONOMYIA (LIPOPHLEPS) LUTEIMARGINATA sp. nov. Plate 3, fig. 39.
Male. — Length, about 2.6 millimeters; wing, 3.3.
Characters as in G. flavomarginata (Brunetti), differing in
details of coloration of the wings and legs.
Thoracic pleura plumbeous-brown, with a single narrow
whitish longitudinal stripe. Legs with the femora brownish
yellow, with a narrow and ill-delimited brown ring just before
46,1 Alexander: Philippine Tipididse, X 33
the tip; tibiae and tarsi dark brown. Wings gray, with a vague
brownish gray pattern, the clearer areas lying chiefly before and
beyond the cord, which is broadly and distinctly seamed with
brownish gray; prearcular and costal regions pale yellowish
white; whitish areas before and beyond stigma; veins very pale
brown, the costal and subcostal veins pale yellow, the cord
darkened. Venation: Scx ending a short distance before the
origin of Rs, this distance about equal to the basal section of
R5 ; anterior branch of Rs straight or very gently sinuous.
Male hypopygium (Plate 3, fig. 39) with the outer dististyle,
od, a gently curved blackened rod, the apex obtuse, near base on
mesal edge produced into a curved black spine, the margin with
conspicuous appressed spines. Inner dististyle, id, a straight
yellow rod, the tip produced into a small blackened recurved
spine. Phallosome, p, terminating in two blackened points, each
produced cephalad into a long black spine.
Mindanao, Davao district, Calian, Lawa, April 24, 1930, at
trap lantern (Clagg) ; holotype, male.
This species agrees very closely with flavomarginata (Bru-
netti) except in the details indicated. Edwards,2 who examined
paratypes of this species, states that all the veins of the wings
are brownish. The Japanese G. (L.) flavocostalis Alexander is
likewise generally similar but differs in all details of the male
hypopygium. The outer dististyle is only weakly spinous along
margin; the inner dististyle is triangular in outline, the outer
end of the triangle being a long pale spine ; phallosome not black-
ened at tips.
GONOMYIA (LIPOPHLEPS) SECRETA sp. n©Y. Plate 1, fi*. 20; Plate 3, fi*. 4<K
General coloration brown ; basal segments of antennae reddish
orange; pleura dark, with a longitudinal, light yellow stripe;
knobs of haltered yellow; legs yellowish brown, without femoral
rings ; wings cream-yellow, with conspicuous pale brown clouds
and washes ; Scj. ending a short distance before the origin of Rs ;
male hypopygium with two dististyles, the outer a powerful
chitinized rod, its tip bifid.
Male. — Length, about 2.5 millimeters; wing, 2.5.
Female. — Length, about 4 millimeters; wing, 3.5.
Rostrum and palpi black. Antennae with the basal segments
reddish orange, the flagellum black. Head pale yellow, the cen-
ter of the vertex darkened.
*Rec. Indian Mus. 26 (1924) 301.
259737 3
34 The Philippine Journal of Science 1931
Pronotum and anterior lateral pretergites light yellow. Me-
sonotal prsescutum brown with a faint grayish bloom; humeral
region restrictedly obscure yellow, the pseudosutural f ovese red-
dish brown ; females with a capillary darker brown median line
on prsescutum ; scutal lobes dark brown, the median area and res-
tricted caudal-lateral angles of the lobes yellow; scutellum
yellow with a conspicuous brown median spot; postnotal medio-
tergite brown, the cephalic-lateral portions more yellowish.
Pleura dark brown, with a longitudinal, light yellow stripe that
is bordered both above and below by scarcely apparent blackish
darkenings. Halteres dusky, the knobs yellow. Legs with the
fore coxae light yellow, the remaining coxae brownish testaceous,
trochanters obscure yellow; remainder of legs pale yellowish
brown, unvariegated, the outer tarsal segments darker brown.
Wings (Plate 1, fig. 20) cream-yellow, with conspicuous pale
brown clouds and washes, including a major area in cell R be-
fore Rs; the cord and outer end of cell 1st M2; conspicuous
longitudinal seams along veins Cu as far as m-cu, cell Cu at base
and along vein 1st A for more than one-half the length; axilla
infumed; stigmal region scarcely darkened; veins pale yellow,
very indistinct, more darkened in the clouded areas. Venation :
SCj ending shortly before the origin of Rs, Sc2 near its tip ; Rs
strongly arcuated; anterior branch of Rs nearly straight; cell
1st M2 closed ; m-cu a short distance before the fork of M.
Abdominal tergites light brown, the anterior-lateral margins
light yellow, the more extensive posterior-lateral margins vel-
vety black; sternites more uniformly darkened; hypopygium
brownish yellow. In female, the tergites blackened, with a res-
tricted yellow area at each cephalic-lateral angle. Male hypopy-
gium (Plate 3, fig. 40) with only two dististyles, the outer, od, a
powerful chitinized rod, the stem straight, the head more en-
larged and bifid, the more slender arm fingerlike, the other arm
flattened, terminating in a comb of microscopic teeth ; inner mar-
gin of stem with a row of powerful fasciculate setae. Inner dis-
tistyle a small pale blade, the tip obtuse. Phallosome, p, com-
plex.
Luzon, Laguna Province, Ube, February 11 to March 3, 1930
(McGregor and Rivera) ; holotype, male; allotype, female; para-
types, 2 females.
Gonomyia (Lipophleps) seer eta by Edward's key to the spe-
cies of the subgenus 3 runs to G. (L.) robinsoni Edwards (Malay
States), a very different fly.
8Journ. Fed. Malay St. Mus. 14 (1928) 104-105.
46,1 Alexander: Philippine Tipvlidse, X 35
GONOMYIA (PTILOSTENA) PUNCTIPENNIS Edwards.
Gonomyia (Ptilostena) punctipennis Edwards, Treubia 7 (1926)
140-141.
A few of both sexes, Lawa, Davao district, Mindanao, taken
at trap lantern, May 5, 1930, by Charles F. Clagg. The species
was described from Buru and will probably be found to be a
widely distributed species in the Malayan and Moluccan islands.
TEUCHOLABIS (TEUCHOLABIS) MAJUSCULA sp. nov. Plate 1, &g. 21; Plate 3,
fig:. 41.
General coloration yellow and black; praescutal stripes con-
fluent; pleura black, striped longitudinally with yellow; knobs
of halteres obscure orange; legs entirely black; wings yellow,
the outer radial cells slightly infumed; male hypopygium with
the outer dististyle a macelike capitate structure.
Male. — Length, about 9 millimeters; wing, 8.
Rostrum nearly as long as remainder of head, black; palpi
black. Antennae black throughout; basal flagellar segments
short-oval, becoming smaller and more elongate outwardly.
Head black, the front and wide anterior vertex sparsely dusted
with gray.
Pronotum very large, yellow. Mesonotal prsescutum chiefly
occupied by three confluent polished black stripes, leaving yellow
areas at the humeri, a transverse median area at the suture and
a tiny spot at each posterior-lateral angle; scutum yellow, each
lobe chiefly covered by polished black centers; scutellum deep
yellow; postnotal mediotergite yellow on cephalic third, the re-
mainder black. Pleura black, with a conspicuous yellow long-
itudinal stripe that extends from behind the fore coxae, passing
beneath the halteres to the abdomen ; dorsopleural region yellow.
Halteres dusky, the knobs obscure orange. Legs with the fore
coxae reddish, the remaining coxae and all trochanters black; re-
mainder of legs entirely black. Wings (Plate 1, fig. 21) with a
strong yellow tinge, the outer radial cells slightly more infumed ;
anterior prearcular cells inf uscated ; veins black. Venation : Sc
long, SCi ending about opposite four-fifths the length of Rs,
Sc2 at near midlength of this vein; Rt in alignment with R1+2J
cell 1st M2 elongate, parallel-sided; m-cu more than its own
length beyond the fork of M.
Abdomen bicolorous, black, the incisures more narrowly
orange, on the tergites this color wider on the caudal margins
than on the bases of the segments. Male hypopygium (Plate 3,
fig. 41) with the tergal region narrowly emarginate medially;
sternite, 9s, convexly rounded, with abundant setae, especially on
36 The Philippine Journal of Science
Bides. Basistyle, b, with the dorsal-apical angle produced into
a black spine; the ventromesal angle with irregular blackened
teeth. Outer dististyle, od, a mace-shaped structure, as figured.
Inner dististyle, id longer, the basal half wider, the distal half
gradually narrower and angularly bent, with three setae at the
angulations, the apex an acute black spine. Phallosome, p, with
a wider dorsal and a narrow ventral plate, both tipped with
long conspicuous setae.
Mindanao, Davao district, Lawa, April 18, 1930, at trap lan-
tern (Clagg) ; holotype, male.
Teucholabis (Teucholabis) majuseula is one of the largest
species of the genus, though exceeded in size by the allied T.
(T.) nigerrima Edwards (Formosa). Both of these species
have Rx in alignment with R1+2, the veins not dipping slightly
caudad at the point of union with R2 as is the case in virtually
all other species of this extensive genus.
TEUCHOLABIS (TEUCHOLABIS) CONFLUENTOIDES sp. nov. Plate 1, fi*. 22; Plate
3, fig*. 42 and 43.
Male. — Length, about 6.5 to 7 millimeters; wing, 6 to 6.5.
Generally similar to T. (T.) confluenta Alexander (Luzon), dif-
fering especially in the structure of the male hypopygium and
the details of venation.
Pronotum extensively pale yellow. Mesonotal praescutum
black, the humeral triangles extensively and conspicuously light
yellow ; scutal lobes blackened, the median region broadly yellow,
crossing the suture onto the praescutum. Dorsopleural region
clearer yellow. Wings (Plate 1, fig. 22) with the pattern band-
ed, much as in confluenta. Venation: Cell 2d M2 much deeper,
exceeding its petiole. Male hypopygium (Plate 3, fig. 42) with
the spine of the basistyle, b, simple. Outer dististyle, od, with
two, or in cases, a minute third, spine, in addition to the long
curved apex. Inner dististyle, id, with a bisetose lobe at base
and on obtuse lobule in addition to the long spinous point.
In confluenta (Plate 3, fig. 43) the spine of the basistyle, 6, is
forked. Outer dististyle, od, a long sinuous rod, with a single
small spine at near midlength. Inner dististyle, id, a simple
black rod.
Mindanao, Davao district, Calian, La Lun Mountains, altitude
5,500 to 5,800 feet, July 2 to 3, 1930 (Clagg) ; holotype, male;
paratypes, 4 males. "Dancing above ferns in semitwilight of
dense mossy forest." — Clagg.
ILLUSTRATIONS
[Legend: a, aideagus ; b, basistyle; d, dististyle; dd, dorsal dististyle; g, gonapophysis ; i, in-
ternal process; id, inner dististyle; od, outer dististyle; p, phallosome; «, 9th sternite;
t, 9th tergite; vd, ventral dististyle.]
Plate 1
Fig. 1. Limonia (Limonia) candidella sp. nov., wing.
2. Limonia (Limonia) latiflava sp. nov., wing.
3. Limonia (Limonia) flavohumeralis sp. nov., wing.
4. Limonia (Limonia) canis sp. nov., wing.
5. Limonia (Rhipidia) luteipleuralis sp. nov., wing.
6. Limonia (Geranomyia) phcenosoma sp. nov., wing.
7. Limonia (Geranomyia) longifimbriata sp. nov., wing.
8. Limonia (Geranomyia) paramanca sp. nov., wing.
9. Limonia (Pseudoglochina) angustapicalis sp. nov., wing.
10. Limonia (Alexandriaria) sollicita sp. nov., wing.
11. Orimarga (Orimarga) rubricolor sp. nov., wing.
12. Epiphragma (Polyphragma) bakeri Alexander, wing.
13. Epiphragma (Polyphragma) parviloba sp. nov., wing.
14. Limnophila (Ephelia) igorota sp. nov., wing.
15. PUaria phcenosoma sp. nov., wing.
16. PUaria carbonipes sp. nov., wing.
17. Gonomyia (Lipophleps) maquilingia sp. nov., wing.
18. Gonomyia (Lipophleps) pallidisignata sp. nov., wing.
19. Gonomyia (Lipophleps) alboannulata sp. nov., wing.
20. Gonomyia (Lipophleps) seer eta sp. nov., wing.
21. Teucholabis (Teucholabis) majuscula sp. nov., wing.
22. Teucholabis (Teucholabis) conftuentoides sp. nov., wing.
Plate 2
Fig. 23. Scamboneura nigrotergata sp. nov., male hypopygium, details.
24. Scamboneura calmnensis sp. nov., male hypopygium, ninth tergite.
25. Scamboneura calianensis sp. nov., male hypopygium, outer dis-
tistyle.
26. Limonia (Limonia) candidella sp. nov., male hypopygium.
27. Limonia (Limonia) flavohumeralis sp. nov., male hypopygium.
28. Limonia (Limonia) canis sp. nov., male hypopygium.
29. Limonia (Geranomyia) phcenosoma sp. nov., male hypopygium.
30. Limonia (Geranomyia) longifimbriata sp. nov., male hypopygium.
31. Limonia (Pseudoglochina) angustapicalis sp. nov., male hypopy-
gium.
32. Epiphragma (Polyphragma) bakeri Alexander, male hypopygium.
33. Epiphragma (Polyphragma) parviloba sp. nov., male hypopygium.
37
38 The Philippine Journal of Science
Plate 3
Fig. 34. Limnophila (Ephelia) igorota sp. nov., male hypopygium, outer
dististyle.
35. Pilaria phcenosoma sp. nov., male hypopygium.
36. Gonomyia (Lipophleps) maquilingia sp. nov., male hypopygium.
37. Gonomyia (Lipophleps) pallidisignata sp. nov., male hypopygium.
38. Gonomyia (Lipophleps) alboannulata sp. nov., male hypopygium.
39. Gonomyia (Lipophleps) luteimarginata sp. nov., male hypopy-
gium.
40. Gonomyia (Lipophleps) secreta sp. nov., male hypopygium.
41. Teucholabis (Teucholabis) majuscula sp. nov., male hypopygium.
42. Teucholabis (Teucholabis) confluentoides sp. nov., male hypopy-
gium.
43. Teucholabis (Teucholabis) confluenta Alexander, male hypopy-
gium.
Alexander: Philippitte Tipulidje, X.]
[Philip. Journ. Sci., 46, No. 1.
PLATE 1.
Alexander: Philippine Tipulid^, X.]
[Philip. Journ. Sci., 46, No. 1.
PLATE 2.
Alexander: Philippine Tipulid^e, X.]
[Philip. Journ. Sci., 46, No. 1.
/T^f^fM^
H
PLATE 3.
ORIGIN OF THE IRRITATING SUBSTANCE IN
MOSQUITO BITE 1
By C. Manajlang
Of the Philippine Health Service, Manila
ONE PLATE
Ludlow2 mentioned the properties of the salivary-gland se-
cretion among the problems concerning malarial infection of
mosquitoes that need further investigation. She said:
As far as I am aware, nobody has yet repeated Schaudinn's observations.
He states that the salivary gland rubbed into an abrasion does not pro-
duce the irritation of mosquito bite, but that, on the contrary, if the oeso-
phageal diverticula be rubbed in, the well known itching effects are exper-
ienced, which he attributes to the enzymes produced by low bacterial forms
in the diverticula. Any facts established about mosquitoes is of value, for
we never know to what practical purposes such knowledge may not be
turned.
Castellani and Chalmers 3 said :
There has been much dispute as to where this substance comes from,
but this appears to have been settled by Schaudinn who triturated the
isolated salivary glands in salt solution which he applied to a wound with
negative results. On the other hand when he applied the isolated oeso-
phageal diverticula to a scratch he obtained the characteristic irritation
and redness. These oesophageal diverticula contain gas bubbles and bac-
teria or moulds. The bubbles were shown by Schaudinn to contain car-
bon dioxide by applying baryta-water to the diverticula, when a precipi-
tate was obtained. The fungi need further investigation, but they or their
products appear to be the real cause of irritation, for when Schaudinn
expressed the carbon dioxide out of the sac the signs characteristic of the
bite were still produced.
While dissecting anopheline mosquitoes for malarial parasites,
I applied isolated glands to five separate needle scratches on
the anterior surface of my forearm. In a few seconds I ex-
perienced itching, followed by the appearance of wheals around
1From the field laboratory of the division of malaria control, Philippine
Health Service, Tungkong Manga, Bulacan.
8 Surgeon General, U. S. Army, Bull. 4 (1913) 90.
8 Manual of Tropical Medicine 3d ed. (1919) 224-225.
39
40 The Philippine Journal of Science 1931
the edges of the scratches, then by a distinct redness of the
surrounding skin in all the scratches. I repeated the test, using
isolated gas-containing diverticula. Of the four trials, one
scratch itched slightly, followed by a small area of redness
around the scratch. The other three scratches did not show
any reaction at all. With results unlike those of Schaudinn's,
I repeated the experiment using more material and two methods
(scratch and prick) of inoculation.
Most of the mosquitoes used were Anopheles ludlowi Theobald
and A. vagus Donitz, which were the dominant species in the
catches at the time. Anopheles maculatus Theobald, A. aconitus
var. filipinse Manalang, Culex quinquefasciatus Say, Aedes
mgypti Linnaeus, a Culex species, four male A. ludlowi, and one
male C. quinquefasciatus were also used.
The scratch method of inoculation seems to have the disad-
vantage of drawing blood, which may dilute or prevent the in-
oculum from penetrating the tissues. Light scratches may not
be sufficient to permit its penetration. They may dry up in
case of any delay in the inoculation. The prick on the other
hand is more like the natural bite. With magnifying-lens con-
trol the inoculum can be picked up at the point of the needle.
This facilitates the entrance of the inoculum at the first prick,
or, if it fails and the inoculum is left on the skin, subsequent
pricks (about twenty) on the same spot will succeed.
METHODS AND RESULTS
Scratch method. — The freshly killed mosquito is dissected in
a drop of normal saline under a magnifying lens. Once the
salivary gland and diverticulum are isolated a series of three
shallow scratches, about 0.5 centimeter long and 2 centimeters
apart, are made with a few strokes of a pointed needle on the
anterior surface of the forearm. This surface is used because
the reaction is clear, the skin being thinner and lighter in
color than that on the posterior surface. With the aid of the
magnifying lens the gland (usually three lobes from one side)
is picked up with the needle and very lightly rubbed several
times into one scratch. Wash the needle in saline, wipe dry, and
repeat the process with the diverticulum (usually the abdominal)
on the other scratch. The third or control scratch is inoculated
with a minute quantity of salt solution from the same drop in
which the mosquito has been dissected. This is done to detect
any soluble substance from any organ which may give a re-
46,1
Manalang: Substance in Mosquito Bite
41
action. The results of the tests by this method are set forth
in Table 1.
Table 1. — Scratch tests.
Organ tested.
Salivary gland (from females)
Salivary gland (from males). _
Diverticulum
Stomach.
Ova _ _ _
Malphigian tubes
Parasites
Con trol
Number
of tests.
30
2
7
4
1
1
1
10
Positive.
30
Slightly
positive.
Nega-
tive.
10
Prick method. — After isolation, the organ to be tested is
picked up on the point of the needle, then inoculated by twenty
light pricks on a fixed point on the skin. I usually select the
root of a hair or the edge of an ink mark. The prick should
be just felt but not deep enough to draw blood. The results by
this method are set forth in Table 2.
Table 2. — Prick tests.
Organ tested.
Salivary gland (from females)
Salivary gland (from males)..
Diverticulum (from females)..
Diverticulum (from males) _ . .
Stomach (females)
Malphigian tubes
Thoracic muscle _
Testes -
(Esophagus .-
Ova - —
Wing
Stomach (from males)
Eye._ _
Parasite
Brain
Control
-Number
of tests.
Positive.
Slightly
positive.
Nega-
tive.
18
18
2
1
1
10
9
1
3
3
2
5
3
6
1
5
1
1
1
2
1
1
1
1
2
1
2
2
2
1
1
1
1
1
1
1
1
12
12
A typical reaction starts with itching immediately followed
by the appearance of a wheal, which enlarges and rises with
increasing paleness in contrast with the spreading redness of
the surrounding skin. The height of reaction is reached at
the moment the wheal begins to lose its pallor and turn red.
A small red indurated area persists at the point of inoculation
42 The Philippine Journal of Science 1931
six to twenty-four hours after a positive test, depending on the
individual's susceptibility. A positive reaction is apparently
more rapid and intense on a perspiring skin than on a dry one.
The reaction is less intense or only slight when the scratch or
puncture is deep and draws blood. It seems, therefore, that
the wheal is due to the entrance of the irritating substance into
the lymphatics in the corium and not in the deeper tissues.
A negative reaction is without itch or wheal. There is only
a redness of the scratch or puncture as the case may be. A
slightly positive reaction produces less itching, a small wheal,
and an area of redness.
All the forty-eight tests with salivary glands from female
mosquitoes (thirty by scratch and eighteen by prick) gave pos-
itive reactions. It will be noted that with the scratch method,
out of seven diverticula tested only one gave a positive reaction
and two slight reactions, while with the prick method the thir-
teen diverticula (from both sexes) all gave typical positive
reactions, except one, which gave a slight reaction. The failure
of most diverticula to react by the scratch method was due to
a deep scratch, to bleeding, to a very shallow scratch, or to a
long interval of time that allowed the serum to dry up between
the time of the scratch and the time the diverticulum was inoc-
ulated. It was often difficult to pick up a diverticulum heavily
loaded with gas bubbles.
OTHER OBSERVATIONS
(a) Salivary glands from five female anophelines were allow-
ed to dry on a slide at room temperature (25 to 35° C.) from
three to five days. Upon inoculation with a little salt solution,
they all produced positive reactions.
(6) Diverticula with gas, or with the gas pressed out, gave
identical reactions. No difference was noted between the reac-
tion of the abdominal diverticulum and that of the thoracic
diverticulum. Diverticulum from the male gave the same re-
action as that from the female.
(c) Fresh or dried salivary gland rubbed into unabraded
skin produced no reaction.
(d) Inoculation by prick of one, two, and three lobes of sal-
ivary gland from one mosquito produced the smallest wheal
with one lobe, and the largest with three lobes. In a repetition
of this test the two-lobe inoculum gave the largest wheal.
(e) Six individuals were each inoculated twice in an identical
manner (prick) with salivary glands from different mosquitoes.
46,1 Manalang: Substance in Mosquito Bite 43
One of them constantly gave a marked reaction with large
wheals and areas of redness, two with lesser, and three with
slight itching, and tiny wheals and transitory redness around
the points of inoculation. Using diverticula, they showed the
same varying degrees of reaction obtained with the salivary
glands.
(/) The reactions obtained from Anopheles, Culex, or Aedes
showed no difference on comparison.
(g) The salivary glands of the male were always much
smaller, slenderer, and more fragile than those of the female.
This probably accounts for the failure to obtain a good positive
reaction for this sex.
(h) The parasites, (Sporozoa or fungus, probably not micros-
poridia) which were not infrequently found in clumps at the
base of the salivary glands, gave a typical bite reaction in one
test and a slight reaction in another. In the fresh state they
appeared as irregular or spherical granular bodies containing
a variable number of very refractile "cysts." Under a high
magnification, these "cysts" were surrounded by colorless, black,
and bluish granules. Stained with Heidenhain's, the parasites
were about the size and appearance of amoebae with vacuoles.
Throughout the tests only salivary glands free from these par-
asites were used. They were not found on the stomach or di-
verticulum.
(i) Positive reactions were observed in a certain number of
stomachs, malphigian tubes, cesophaguses, ova, testes, etc.
SUMMARY AND CONCLUSIONS
1. The salivary gland of the mosquito inoculated into the
skin produced a typical bite reaction, contrary to Sehaudinn's
finding.
2. Sehaudinn's reaction using the diverticulum was confirmed
by the prick method in ten tests with the diverticula from the
female mosquito and in three tests with those from the male.
3. The irritation of mosquito bite must, therefore, be due to
injection of the salivary-gland secretion or diverticular con-
tents, or both, and not to diverticular origin only.
4. Typical bite reactions were also obtained from parasites
(Sporozoa or fungus) and from the other organs; such as,
stomach, oesophagus, testes, and ova.
5. Different degrees of susceptibility to the bite were tested
and demonstrated in six individuals.
ILLUSTRATIONS
[Microphotographs by C. M. Urbino, Philippine Health SerTice.]
Plate 1
Fig. 1. a, Thoracic diverticulum with gas; b, abdominal diverticulum with
gas; c, stomach; df Malphigian tubes; e, two detached lobes of
the salivary gland; about X 55.
2. Salivary gland from A. vagus Donitz with eight lobes and a cluster
of parasites at the base; about X 160.
45
MANAiiANt;: Substance in Mosqi'ito Bite,]
[Philip. Journ. Scf.» 46, No. 1.
PLATE 1.
MALARIA TRANSMISSION IN THE PHILIPPINES, III *
DENSITY AND INFECTIVE DENSITY OF ANOPHELES FUNESTUS
GILES
By C. Manaiang
Of the Philippine Health Service, Manila
The present paper is a continuation of the preceding two
articles and is based on certain data compiled from observa-
tions that began in September, 1927, in La Mesa camp of the
Novaliches water project and extended during the following
two years to seven other camps and two barrios far from, and
independent of, the project. An attempt will be made to show
the significance of density in the transmission of malaria without
considering the suitable human-carrier factors. About 22,000
Anopheles funestus Giles were caught, mostly by trapping,
and were dissected. With these were 298 infected mosquitoes
found in six out of ten places, as follows: In South Portal
camp 218 positive mosquitoes were caught; in North Portal, 26;
in La Mesa, 32; in Atlantic, Gulf, and Pacific Company camp,
13 ; in Tungkong Manga, 7 ; and in San Francisco del Monte, 2
(oocysts only). No positive mosquitoes were caught in Nova-
liches Barrio, Bigti, Santo Cristo, or Alinsangan camps.
The traps used were of standard dimensions and the time
employed by the catcher was practically the same in all areas.
Approximately the same number of hours was also used in catch-
ing mosquitoes without traps. The densities obtained through
these collections are only approximate, but the duration and
regularity of observation are considered sufficiently adequate
to counterbalance the errors.
The appended tables give the catches of A. funestus in all
areas with the results of dissection, densities for the positive
and negative months, and densities in four negative places.
Infective (sporozoite rate) densities are expressed as one for
every so many mosquitoes dissected by individual months and
as monthly averages during each year. Table 7 is a summary
1 From the field laboratory, division of malaria control, Philippine Health
Service, Tungkong Manga, Bulacan.
47
48
The Philippine Journal of Science
1931
of findings in five places, by years, where sporozoites were dis-
covered, giving the average monthly and infective densities by
years, and the average densities for negative months, if any,
to compare with the average densities of positive months. Ta-
ble 6 shows the catches in four areas where no infected funesttis
were found.
Table 1. — Catches and findings in South Portal.
Year and month.
Number
caught.
Positives.
Infective
(sporo-
zoite)
density.
One in —
Stomach.
Salivary
gland.
Total
infected.
1927
234
723
201
567
498
562
469
735
490
371
671
255
223
6
9
3
7
3
16
6
23
16
8
9
3
4
2
4
3
6
8
17
11
10
7
7
2
2
6
8
13
6
13
11
33
17
33
23
15
11
5
10
177
181
67
95
62
33
43
73
70
53
335
127
37
1928
February. ..
May. . - -
July __ _
Total _ -
5,765
107
83
190
Average catch per positive month (density), 1928_
Average monthly i nf ective density
480
•69
93
23
170
62
1929
January -. - - - .-
93
65
90
170
124
313
148
81
43
29
47
57
2
3
1
2
2
i
4
1
2
1
2
7
2
4
2
April
May.__
July -
1
1
October
November .
1
1
--^
«>993
124
67
12
8
20
Average catch per positive month (density), 1929 _
Average catch per negative month ( density)
Average monthly infective density
» 124
* Computed on catches during positive (stomach and salivary gland) months only.
b Positive months only. December, 1927, to February, 1928, inclusive, by exposure.
Trap used since March, 1928.
46,1 Manalang: Malaria Transmission 49
COMMENTS
South Portal (Table 1) was by far the most malarious (more
transmission) of the six places, judging by the mosquito in-
fection. In this camp, the year 1928 was more malarious than
1929, there having been not only more funestus but fewer bites
were necessary for infection. The average monthly density
and the infective density were 480, and 1 in 69, respectively,
in 1928, as compared with 124, and 1 in 124, respectively, in
1929. Within the average twenty-three catching days per
month, the susceptible human bait in the trap had seven chances
monthly of becoming infected if all the mosquitoes bit him, or
an average of almost one infective bite every three days during
1928, In 1929, his chance was one infective bite in twenty-three
days; in December, 1927, two infective bites in twenty-three
days.
At North Portal (Table 2) with its monthly density of 412,
and infective density of 1 in 515, in 1928, it would have required
the bites of all the mosquitoes coming into the trap for twenty-
nine days to assure an infective one being included, while in
1929, with 833, and 1 in 1,250, as monthly and infective densities,
respectively, it would have required thirty-five days before in-
fection could have been received.
La Mesa (Table 3) was more malarious in 1928 than in 1927,
in spite of a larger number (more than three times) of funestus
in 1927. The infective density of 1 in 400 and the monthly
density of 500 in 1927 would only give a little over one infective
bite during twenty-three catching days, while the lower density
of 120 in 1928 would give more than two infective bites (the
infective density being 1 in 52) . The duration of observations
in the other places does not permit comparison of infection of
one year with another, but a comparison of one place with an-
other shows again the role of density in the amount of malaria
transmission.
In 1929, the average density during the positive months (Jan-
uary to May) in Tungkong Manga was 846 for a month of
twenty-three catching days. The infective density was 1 in
1,411, so that it would have required an exposure of thirty-nine
days before an infective bite could have been received. This
is in accord with James's 2 statement that "numerous anophelines
2 Malaria at Home and Abroad. John Bale, Sons and Danielsson, Ltd.,
London (1920) 13.
259737 4
50 The Philippine Journal of Science
Table 2. — Catches and findings in North Portal.
1931
Year and month.
Number
caught.
Positives.
Infective
(sporo-
zoite)
density.
One in—
Stomach.
Salivary
gland.
Total
infected.
1927
8
30
105
0
0
7
0
0
0
110
989
209
771
1
3
1
3
1928
January. __.«_»-.- - - - - - - - -
April
July
*
12
1
1
1
2
1
1
14
1
2
110
469
771
Total - -
•2,059
17
4
21
Average catch per positive month (density)
Average catch per negative month (density)
412
56
b515
1,807
572
1929
1,307
620
572
366
68
46
26
10
5
1
1
1
1
2
1
1
February - - - - -
April
Total - -
•2,499
2
2
4
Average catch per positive month (density)
Average monthly infective density
833
87
^1 ,250
• Positive months only.
b Based on total catches in positive months. Trap used since September, 1928.
of a species which is an efficient carrier are associated with
little or no malaria," and with Gill's 8 reference to "anopheles
sine malaria, one instance of which is the large measure of
'control,' achieved over malaria in Italy by the method of 'boni-
fication' in spite of the fact that this measure has actually led,
in some instances, to increased prevalence of anophelines." Had
the density in Tungkong Manga prevailed in La Mesa (com-
8 Trans. 7th Cong. Far Eastern Assoc. Trop. Med. 2 (1927) 630.
46,1
Manaiang: Malaria Transmission
Table 3. — Catches and findings in La Mesa.
51
Year and month.
Number
caught.
Positives.
Infective
(sporo-
zoite)
density.
One in —
Stomach.
Salivary
gland.
Total
infected.
1927
September
October - . - .._---
669
473
512
346
3
4
6
1
4
1
3
8
6
669
128
November . -
December .--»_-._ - -
Total
2,000
500
13
5
18
Average catch per month „ „ . .
Average monthly infective density
400
39
1928
January
116
61
227
0
7
0
0
0
18
1
3
4
February. _
March
5
4
9
57
April.. - -
May - -
June. »-- - -
July .... . .
August .- ...
September _. - -
1
1
Total _
•361
120
34
52
7
7
14
Average catch per positive month (density)
Average monthly infective density
1
a Positive months only. Catches by exposure.
pare with May, 1928, South Portal) during January, February,
March, and April, 1928, with an infective density of 1 in 52,
the rate would have meant sixteen infections in twenty-three
days; in which case only a little over one day of exposure
would have been required to cause one infection. This would
have resulted in an explosive epidemic that would probably have
disappeared rapidly from May till October (average density
6 per month). Conversely, had the monthly infective density
of Tungkong Manga, 1 in 1,411, occurred in La Mesa, with its
monthly density of 120, it would have required two hundred
eighty-one days of exposure before a positive bite could have
been contracted, so that there would have been no malaria at
all in the locality (compare with 1929, Tungkong Manga) .
The fluctuation of individual monthly density and the cor-
responding infective number (the latter due to the number of
suitable human carriers, whose movements and accessibility to
the vector could not possibly be under control) is interesting
52 The Philippine Journal of Science 1931
Table 4. — Catches and findings in San Fmncisco del Monte.
Year and month.
1928
June
July..
August
September-
October
November. .
December-.
Total,
Average catch per positive month (density). .
Average catch per negative month (density) _
1929
January
February
March
April
May
June
July ---
August....
Total-
Average monthly catch (density).
Number
caught.
Stomach.
2
1
5
32
108
69
80
"108
108
108
29
4
8
151
25
Positives.
Salivary
gland.
Total
infected,
a Positive month only (October). Trap used from September, 1928.
and explains how easily a person may contract the disease
during a very short visit to a malarious place. Take, for ex-
ample, South Portal in May, 1928, when a trap density of 562
with an infective density of 1 in 33 was capable of seventeen
infective bites in twenty-three catching days. During the
month, one could easily have contracted the disease in a little
over one night's sleep in the camp. Yet, in terms of the in-
fection rate, the usual way of expressing malaria in numbers
of mosquitoes, only 3 per cent of them had sporozoites in their
salivary glands capable, at the time, of transmitting the disease;
certainly a low figure by itself if one is to receive only one bite.
But since, in nature, an exposed individual receives several or
many bites during a single night, his chances of infection must
rise in proportion to the number of bites he receives. To pro-
duce an infection for example, thirty-three individual mosquito
bites would not be too many or too noticeable a number to be
received in, say, two nights during slumber. The percentage
46,1 Manalang: Malaria Transmission 53
method of expressing malaria in mosquitoes seems to have led
experienced students to explain the very high incidence of ma-
laria in man by assuming that the vector bites only in the house,
and that once infected it stays in the same house, or that if it
leaves it usually returns, or by the findings (experimental or
epidemiological) that an infective anopheline can infect several
people in one night or many in several consecutive nights
(James) .4 A similar argument was put forth by Swellengrebel 5
of the Malaria Commission of the League of Nations in the
discussion of the subject, "Where does A. maculipennis infect
man?" to prove that it (maculipennis) bites in the house and
not in the open. He says,
Even under extremely favourable conditions, the number of infected
A. maculipennis is so small that the chances of being infected by anopheles
in the open seem to be infinitesimal, even if this insect were in the habit
of biting in the open, which is generally supposed not to be the case.
On the other hand, if we suppose that at least a portion of the anophe-
lines regularly visiting houses remain there sufficiently long to become
infective after having become infected with malaria parasites, this would
explain not only human infections, even with a small parasite index
among the anophelines but also the house infection so often observed.
This house infection can hardly be explained by random infection.
. . . His figures showing the greater prevalence of infective
mosquitoes in the houses as compared with the stables to prove
his contention are as follows :
In winter (Sella in Fumicino: October-December, 1918, 3.8-4.6 per
cent infected in houses, 0.49-0.85 per cent in stables) ; (Swellengrebel,
Amsterdam, October- December, 1920, 4.99 per cent infected in houses,
0.66 per cent in stables). But in summer this may be different (Sella,
June-September, 1919, 0.5-3.3 per cent in houses; 0.27-2". 00 per cent in
stables).
These figures are not significant, unless it is possible to show
that the stable rates of infection were based only on the numbers
of maculipennis actually concerned in the malaria transmission
in man, but resting in the stable, and that the rates were not in-
fluenced by the numerous anophelines in the stable that fed
solely on animals. A corrective factor based on precipitin test
should have been mentioned. King 6 observed that a high mos-
quito density (575 bites during the season) offsets a low infec-
4 See reference, footnote 2.
6 Report on its tour of investigation in certain European countries in
1924. Geneva (March, 1925) 178.
6 Southern Med. Journ. 17 (1924) 596-597.
54 The Philippine Journal of Science 1931
tion rate (0.107 per cent) and accounts for the prevalence of
malaria. It is believed that a study of the densities and in-
fective densities in different sections or in houses of a locality-
over an adequate period of time will explain house infection
more satisfactorily, as only then can the theory of many human
infections from a single mosquito (a rather difficult task to
demonstrate directly in nature, particularly if the vector is wild,
although understandable epidemiologically) be proved with reas-
onable scientific certainty. In his summary (p. 183) , however,
Swellengrebel said:
The evidence that man is infected outside his habitations (in the wider
sense of the word) is still insufficient, but strong enough to warrant further
enquiry.
He mentioned that during their visit to Fumicino (p. 180)
Grassi emphasized the discordance between the high malaria
incidence and the low number of anophelines in the house and
set forth the following practical question:
If the number of anophelines in the immediate surrounding of man
has so little to do with the incidence of malaria, is this not an indication
that if once anophelines are present in a certain minimum quantity,
other factors influencing the incidence of malaria tend to become of
such predominating importance that it matters little whether the initial
number of anophelines is maintained or multiplied tenfold? If this, or
anything like it, were true, any attempt to diminish the malaria-inci-
dence by reduction of the number of anophelines will be useless, unless
the reduction reaches this hypothetical minimum.
The last question is in accord with Ross 7 and explains the
excellent results of Watson's antilarval measures in many of his
projects in Malaya, the water-cistern control in Palestine, par-
ticularly Jerusalem, mosquito control in the United States, and
Hackett's projects in Italy. It is also clear that an initial hy-
pothetical anopheline density and infective density in one locality
capable of inflicting an infective bite every two or three days
(compare with May, 1928, South Portal) can be maintained or
multiplied tenfold and still infect 100 per cent of the exposed
and susceptible individuals within a short period of time because
the number of new cases will appear rapidly and, therefore,
outnumber the recoveries. But a density and infective density
of one infective bite in one month as an initial minimum hypo-
T "That if the number of malaria-bearing Anophelines is below a certain
figure that limit (fixed limit of malaria) is zero." — The Prevention of Ma-
laria (1911) Sec. 28.
46,1 Manalang: Malaria Transmission 55
thetical number in another locality (Tungkong Manga, 1929),
if only maintained, cannot possibly infect a large percentage of
the people within a short time because the number of new cases
are too few and far apart to outnumber the recoveries. To in-
fect 100 per cent it would be necessary to multiply the initial
density many times over.
The summaries given at the end of the present paper are
believed sufficient answer to the practical question quoted from
Swellengrebel. The hypothetical number must be, according
to the data presented, not only variable in the same place at
different periods, but must also differ in different localities,
which accounts for the successes and failures of the mosquito-
control measures that have always been the cause of misunder-
standing between the Malaria Commission and the antimosquito
enthusiasts, as evidenced in their conclusion written by Swel-
lengrebel8 (p. 189) as follows:
Although during our tour we have seen many instances of anti-larval or
anti-adult measures, there was not one of them in which the efficiency of
the measure had been proved by its influence on the prevalence of malaria.
This does not mean that there had not been such an influence, although in
many instances this was probably so, but that the methods to collect
the necessary statistical material had been inadequate.
On page 174 he says:
We have tried to form a judgment on the effect of the anti-larval
measures demonstrated to us, not by the reduction of the malaria rate
(because (1) this reduction usually could not be demonstrated, owing to
the absence of reliable statistical material; (2) it often coincided with
a reduction in other places where no such measures Were taken; (3)
this measure was never taken without others, notably quininisations, the
effect of the two being difficult to distinguish) but by the prevalence of
larvae in the breeding places and of adult anophelines in houses especially
in stables.
In this instance the conclusion is unavoidable that the Ma-
laria Commission of the League of Nations erred in undertaking
to form a judgment (on the influence of antilarval measures in
particular places where malaria has been reduced) based on
the prevalence of larvee in the breeding places and on adults
in houses and stables at the time of their visit, when data on
the prevalence of both larvae and adults (which should have
been measured by the same standard used by the commission)
before the control was instituted, were either not available or
utilized for the purpose of comparison.
8 See reference, footnote 5.
56
The Philippine Journal of Science
1931
Table 5. — Catches and findings in Atlantic, Gulf, and Pacific Company
and Tungkong Manga,
ATLANTIC, GULF, AND PACIFIC COMPANY.
Number
caught.
Positives.
Infective
(sporo-
zoite)
density.
One in —
Year and month.
Stomach.
Salivary
gland.
Total
infected.
1928
311
29
12
7
7
3
3
10
3
104
April .-_-
Total
•340
10
3
13
Average catch per positive month (density)
Average catch per negative month (density)
170
10
113
TUNGKONG MANGA.
1928
December.
1929
January
February
March
April
May _-
June
July
August.
September... .
October
November
December
Total.
Average catch per positive month (density)..
Average catch per negative month (density) -
Average monthly infective density
273
1,367
849
917
1,086
13
39
18
4
1
20
166
282
» 4,232
846
100
849
1,086
13
1,411
a Total for positive months only.
Manga since June, 1929.
A, G, & P. Co. by exposure. Trap used in Tungkong
The unaccountable marked drop in A. funestus densities in
1929 in North Portal and Tungkong Manga (February in the
former and May in the latter), two distant places (Tables 3 and
5), is significant and explains the sudden diminution or disap-
pearance of transmission in certain localities, which are often
attributed to control measures. It would not be surprising if
some of the places investigated by the commission were instances
46ri Manalang: Malaria Transmission 57
of this nature. On the other hand, the high density maintained
in South Portal in 1928 is most favorable for continuous or
hyperendemic malaria which, as has been experienced, was not
amenable to larval control, while the monthly and infective den-
sities in 1929 (124, and 1 in 124, respectively) might have easily
been affected by larval destruction.
It is also clear how a small number of A. funestus (120 per
month of twenty-three catching days in La Mesa in 1928 or
5 per night) can produce considerable malaria (infective den-
sity, 1 in 52). A similar condition might have prevailed at
Ennur as cited by James,8 as it would have required three ex-
perienced observers four days to catch the sixty mosquitoes.
The data presented show clearly that a quantitative measure
alone of the transmitting species, no matter how careful and
systematic it is, does not give the same amount of necessary
information that their systematic collection and examination
for sporozoites do on the epidemiology of malaria, when it is
considered that the factors of suitable human carriers and sus-
ceptibles are continuously variable. In case the vectors bite
animals with frequency, the density should be corrected by
using a precipitin-test factor. Mosquito density in a locality at
a given period is of importance only when the infective number
in the same locality during that period is known. With these
two adequate data available, the amount of malaria in a sus-
ceptible community is directly proportional to the adult density
of the carrying species. The procedure for these determina-
tions seems simpler, and the results scientifically more accurate
and direct because only mosquitoes are dealt with, dispensing
with the uncontrolled human-carrier movement and the still
more difficult work of determining who and how many are the
carriers and for how long a period they will be suitable mos-
quito infectors, and the equally difficult task of differentiating
between the new attack and the relapse in the measurement of
malaria in man.
If the catches in the four negative places (Table 6) could
be considered sufficient, the low density and probably a 1 to
a very high figure of infective density (for example, 1 in 2,000
or more) explain why evidence of malaria transmission was not
8 "Thus at Ennur in Madras where most of the inhabitants suffered
from malaria, the infecting species of anopheline was so rare that three
experienced observers were occupied for several days in catching about
sixty specimens." See reference, footnote 2.
58
The Philippine Journal of Science
1931
observed and new cases were not as prevalent in them as in the
other camps with similar topography and mosquito fauna.
Table 6. — Catches by exposure in four negative places.
Place.
Year and month.
Number
caught.
No valiches
December. .
January
1927
2
122
150
274
89
Do _
1928
Do -
February
Do _
March .
Do _-- -
April .„.. -
Total _-_ _
585
Average per month (density)
117
Santo Cristo_ ... - . -- ---
May
1929
13
11
16
0
1
2
100
123
Do - -
June ...
Do -
July
Do _.
August -
Do _
September..
October
Do
Do - -
November..
December..
Do
TotaL _ -
266
Average per month (density) - „ _ _ «
33
Bigti -
October
1929
2
107
82
November..
December. _
Do
191
64
November.-
December. .
1929
23
18
Do
41
Average per month (density)
20
SUMMARY
1. Data collected from ten places consisting of dissections of
about 22,000 Anopheles funestus from September, 1927, to De-
cember, 1929, inclusive, show monthly (twenty-three catching
days) and yearly variations in densities in different localities
and in different periods of the same locality as measured by
systematic catches by trapping with human bait or by exposure.
46,1
Manalang: Malaria Transmission
59
Table 7. — Summary showing average monthly densities, positive and ne-
gative months, and infective numbers in five places where sporozoites
were found by dissection.
Place.
Year.
Monthly
density
positive
months.
Monthly
density
negative
months.
Average
monthly
infective
number.
One in —
South Portal _ _
1927
1928
1929
1927
1928
1928
1929
1928
1929
234
480
124
600
120
412
833
170
846
177
69
124
400
52
515
1,250
113
1,411
Do _ _ _
Do
67
La Mesa
Do _
34
56
87
10
75
North Portal
Do_ _
Atlantic, Gulf, and Pacific Company
Tungkong Manga
2. The infective density or number (sporozoite rate) varied
in different places, and in the same place in different periods due
to variations in the ever changing human-carrier factor.
3. Much malaria with few* transmitters and vice versa exist
and can be explained by a knowledge of the density and the in-
fective number of the place at the time.
4. Natural unexplained marked decline in density has been
observed in two places and explains the sudden disappearance of
transmission in certain uncontrolled localities, which are often
attributed to antimalarial measures in controlled areas.
5. The numerical prevalence of the transmitting species alone
means little in the epidemiology of malaria, neither can a known
density in one locality be utilized for comparative purposes in
another. However, the direct relationship of the vector's den-
sity to malaria transmission in a locality at a certain period,
when the corresponding infective number for that locality and
period is known, has been shown to operate in nature.
6. A study of the densities and sporozoite rates of the trans-
mitting species in several localities of the Novaliches water
project over an extended period has revealed at least some of
the fundamental causes of the different behaviors of malaria in-
cidence that were formerly obscure.
7. Important documents have been quoted and discussed in
the light of the present findings. They show that opinions on
numerical anopheline prevalence and malaria incidence have
apparently been based on inadequate field data.
FRESH-WATER SPONGES OF THE PHILIPPINE ISLANDS
By N. Gist Gee
Of the Rockefeller Foundation, Peking, China
FOUR TEXT FIGURES
Very little has as yet been done toward the study of the fresh-
water sponges of the Philippine Islands. Doubtless, a little
time spent by collectors in examining the lakes, ponds, and
streams of the Islands would yield a rich supply of interesting
new materials. So far as I can learn only four species of this
group have up to date been recorded from the Islands and all
of them were described as new to science.
The first species is recorded by W. Weltner,1 who describes
Ephydatia fortis as a new sponge and gives the following note
concerning its occurrence and collector: "Libmananfluss auf
Luzon, Museum Berlin, Jagor leg." The type specimen of this
sponge is in the Berlin Museum.
The other species were described by Annandale from material
in the United States National Museum. He 2 described two new
species, Spongilla philippinensis and Spongilla dementis, which
were collected by Mary Strong Clemens in January, 1907, at
Camp Keithley, Lake Lanao, Mindanao, at an elevation of 2,250
feet. In another paper Annandale 3 again reports upon Philip-
pine fresh-water sponges. Paul Bartsch collected some speci-
mens of fresh-water sponges from Vicars Landing, Lake Lanao,
Mindanao, in May, 1908. These were determined by Annan-
dale to be S>. philippinensis; this makes a second locality in Lake
Lanao in which this sponge has been found.
Other fresh-water sponges were collected by Hugh M. Smith,
of the expedition of the Bureau of Fisheries steamer Albatross,
December 26, 1907, from Taal Lake, on the east side of Taal
Island, Luzon. Annandale called this sponge a new species and
named it Spongilla microsclerifera.
1 Spongillidenstudien III, Archiv fur Naturg. 1 (1895) 141.
2 Proc. U. S. Nat. Mus. 36 (1909) 629-632.
8Proc. TJ. S. Nat. Mus. 37 (1909) 131, 132.
61
62 The Philippine Journal of Science 1931
The type specimens of the last three sponges are in the United
States National Museum.
Without doubt there are other fresh-water sponges to be
found in the Philippine Islands and the writer would be pleased
to receive specimens for study. They may be dried in the
shade, wrapped m soft paper, and then sent by post in small
tin boxes or in strong, light, wooden cigar boxes. Specimens
should, if possible, contain gemmules, but even when without
them they should be gathered. A former correspondent, now
deceased, wrote of the great abundance of these sponges on the
fishing traps in the waters near Los Bafios, but he unfortun-
ately did not send me specimens of these.
SPONGILLA PHILIPPINENSIS Annandale, 1909. Text fi*. 1.
Historical statement — This sponge was collected in January,
1907, by Mary Strong Clemens, at Camp Keithley, Lake Lanao,
Mindanao, Philippine Islands, at an altitude of 2,250 feet. It
was sent by the United States National Museum, together with
other sponges, to Dr. N. Annandale, of the Calcutta Museum,
for identification. His illustrated description was published in
1909.4 Since the very small bit of this sponge, which the
United States National Museum has kindly given me, contains
no gemmules and is too small to give any idea of the structure
of the sponge, I shall quote in full Annandale's original detailed
description.
Habitat. — Paul Bartsch collected additional specimens of this
species at Vicars Landing in the same lake in May, 1908. They
were taken in shallow water and were attached to the submerged
drift around the edge of the lake.
General characteristics. — "The sponge has evidently formed a
sheet of considerable size adherent to some solid body, but has
been broken into small pieces in the type specimens, which are
about one centimeter thick. The surface is smooth with nu-
merous oscula level with it. There is no trace of branches."
Color. — "Externally the sponge appears to have been bright
green in color, but the basal parts are yellowish." It is gray in
alcohol.
Structure. — "The texture is light and friable, by no means
elastic. In vertical section both radiating and transverse fibers
are visible to the naked eye and the sponge has a distinctly
reticulate appearance, although the vertical interspaces are
4Proc. TJ. S. Nat. Mus. 36 (1909) 629-631.
46,1
Gee: Fresh-water Sponges
63
«
a
Fig. 1. SpongUla philippvnensis Annandale. o, Showing both the smooth and the spinet*
skeleton spicules ; also one gemmule spicule (after Annandale) ; b, smooth skeleton
spicules with axial canals clearly showing. (Drawing by C. F. Wu.)
much more conspicuous than the horizontal ones. Wide cir-
cular canals penetrate the sponge in a course parallel to the base.
Comparatively little spongin is present. Under the microscope
it is evident that the radiating fibers are much more coherent
and regular than the transverse ones. On the external surface
of the sponge a network of the horizontal spicules can be distin-
guished. There is a delicate, basal structureless membrane.
The ectodermal membrane has perished."
Skeleton spicules.— "The skeleton spicules measure 0.174 mm.
to 0.278 mm. in length and on an average 0.021 mm. in greatest
transverse diameter. They are very sharply pointed at both
ends, straight or nearly so, smooth or somewhat sparsely covered
with extremely minute projections, the ends being always
smooth."
I find these spicules to range between 229 and 271 p in length
and between 14 to 20 /* in diameter. In the slides that I have
examined I have found no spined spicules at all.
Flesh spicules. — There are no flesh spicules.
Gemmules. — "There are few gemmules, those that exist oc-
curring singly in the substance of the sponge and being free.
They have a blackish color, are spherical, measuring on an aver-
age 0.609 mm. in diameter. Each is provided with a single
64 The Philippine Journal of Science 1931
aperture, to which a short, straight, rather stout foraminal tu-
bule is attached. The inner chitinous coat is rather thin, but
the granular coat is well developed and contains many spicules,
which are arranged horizontally or nearly so as a rule, but
sometimes to a slight extent tangentially."
Gemmule spicules. — "The gemmule spicules are very variable
in length, measuring from 0.0798 mm. to 0.122 mm. in length
and about 0.0031 mm. in transverse diameter. They are cylin-
drical, straight or nearly so, armed with somewhat irregular
spines, which are often slightly retroverted at the two ends.
Sometimes there is a single straight spine at either end, but
often the spicule ends abruptly and is surrounded by a ring of
spines in such a way as to suggest a rudimentary rotule."
Type. — The type is preserved in the collection of the United
States National Museum in Washington. I have a small, gem-
muleless specimen (No. 54337) from that museum in my collec-
tion.
Distribution. — Spongilla philippinensis has so far been found
only in Lake Lanao, but a closely related form, S. sceptrioides,
has been described from New South Wales and Queensland,
Australia.
Remarks. — From the descriptions and illustrations of S. phil-
ippinensis and S. sceptrioides given by Annandale, it seems that
the spicules of these two sponges are very similar and I doubt
very much that they are both entitled to separate specific rank.
It is very desirable that new material of both of these sponges
be collected for further comparison before a final decision is
reached. S. philippinensis is also related to S. alba but is read-
ily distinguished from it "by having minutely spined megas-
cleres, green corpuscles, slender gemmule spicules with short
spines and no free microscleres."
Concerning the specimens collected by Bartsch in May, 1908,
Annandale writes that while no gemmules were present, the
sponges were full of embryos. "The embryos lie in the inter-
stices of the skeleton and have no protecting membrane as is
the case in some oriental species (Records of Indian Mus., Vol.
1, p. 269 (1907). They are so numerous that in preparations
made by boiling pieces of the sponge in nitric acid their mi-
nute immature skeleton spicules are present in sufficient num-
bers to appear to be a feature of the species and might easily
be mistaken for free microscleres. True flesh spicules are,
however, absent."
46fi Gee: Fresh-water Sponges 65
SPONGILLA CLEMENTIS Annandale, 1909. Text fig. 2.
Historical statement. — In January, 1907, Mary Strong Cle-
mens collected this species at Camp Keithley, Lake Lanao, Min-
danao, Philippine Islands. The altitude of the lake is 2,250
feet. This sponge was described and illustrated by Annan-
dale.5 Annandale described a similar sponge from Tali Fu,
Yunnan, China, calling it at first Spongilla yunnanensis 6 but
later,7 he corrected this and designated that sponge also as
S. dementis. The same author, in 1916, also described the same
species from Lake Biwa, Japan.8 The description that follows
is based upon Annandale's descriptions of the several forms
examined by him and my observations upon a small specimen
of the original Philippine material kindly furnished me by the
United States National Museum, a small bit of the Tali Fu
sponge kindly provided by the Indian Museum, and a splendid
series of specimens from Japan kindly given me by Doctor
Kawamura, of the Biological Station at Otsu.
Habitat. — In Lake Lanao this sponge was found growing in
close association with Spongilla philippinensis. In Yunnan it
was found growing on small stones in the lake where it formed
small rounded masses. In Lake Biwa it grew in three quite
distinct phases: (1) It formed flat crusts of irregular outline
usually less than 10 millimeters in thickness. (2) This phase
also formed crusts, but was more massive than the first and at
times developed "thick ramifying horizontal branches." Both
of these phases were found growing on the pillars of bridges
and piers and other similar supports, and sometimes covered
considerable areas. (3) The third phase differs decidedly from
the other two; it formed "compact, ovoid, spherical, irregularly
massive or pedunculate masses." These masses grew on cer-
tain living mollusks or were found on stones, sticks, or lying
free on the clean, sandy bottom of the lake in deep water.
General characteristics. — "In general appearance and color,
this sponge, judging from the dry specimens, closely resem-
bles Spongilla philippinensis, but the surface is usually covered
with a network of deep, broad furrows which separate small
elevated areas of more or less circular form. The oscula oc-
cur on these elevated areas and are large and numerous. Prob-
6Proc. U. S. Nat Mus. 36 (1909) 631-632, fig. 4.
• Rec. Indian Mus. 5 (1910) 197.
TMem. Asiatic Soc. Bengal. 6 (1918) 201.
8Journ. Coll. Sci. Tokyo 39 (1916) 7.
259737 5
66
The Philippine Journal of Science
1931
& ££2
Fig. 2. SpongiUa dementis Annandale. a, Spined and smooth skeleton spicules from
specimens from Lake Biwa, Japan; the smaller spicules show the variations in gemmule
spicules (after Annandale) ; b, skeleton and gemmule spicules of specimen from Lake
Lanao, Mindanao; only smooth skeleton spicules were found (after Annandale).
ably in the fresh sponge the furrows are roofed in by the ec-
todermal membrane/'
Color. — The color of the Philippine specimen of S. dementis
was quite similar to that of S. philippinensis, green on the sur-
face and yellowish underneath. The China representative of
46»1 Gee: Fresh-water Sponges gf
this species was of a dull greenish color when growing. An-
nandale describes the colors of the various specimens of this
sponge from Japan as "leaf green, grayish or yellowish, . . .
tinged with green, but the color never extends to the inte-
rior, . . . grayish or whitish." The majority of our specimens
(dry) from Japan are a light brown or straw color; a few are
of a grayish color.
Structure.— "In vertical section the transverse fibers of the
skeleton of this species from the Philippine Islands are seen
to be stouter and more regular than those of S. philippinensis,
being hardly inferior to the radiating fibers in these respects!
so that the skeleton forms a much more regular network than
is the case in the other sponge."
There is a good deal of variation in the consistency of the
Japan sponges. Some of them have transverse fibers strong
enough to make them quite firm and hard, while others are
soft and easily crumbled. In the Philippine representatives
of this species "there is a stout chitinous membrane, which sends
bunches of hollow, root-like processes downwards at intervals.
These do not appear to be in any way connected with the pri-
mary skeleton fibers. There are numerous scattered skeleton
spicules in the basal membrane/'
In the Japan forms, the oscula are usually numerous, large,
and round and open upon the smooth surface of the sponge.
Slight elevations or ridges may sometimes be formed around
the oscula, or in some phases they may even develop crater-
like cones.
Skeleton spicules. — Here again it is difficult to cover all var-
iations in skeleton spicules in one general description. My spec-
imens of both the Tali Fu and the Lake Biwa sponges have
only smooth spicules in their skeletons. The Philippine sponge
has a large majority of its spicules with spines, although the
number of spines may vary from a very few small ones, which
appear to be only granulations, to a rare spicule now and then
that is thickly studded all over, except at the tips, which has
prominent spines (fig. 3).
The spicules of the Japan forms closely resemble those of the
Yunnan specimen, and I am of the opinion that these two species
more closely resemble the form described by Annandale as
S. dementis, even though I evidently have a bit of what seems
to be the original Philippine material from the United States
68 The Philippine Journal of Science 1931
National Museum from which Annandale described this spe-
cies. The resemblance between this Philippine sponge, S. de-
mentis, and the one which Annandale describes as S. sceptrioides
in the same paper is quite close. I think that additional ma-
terial bearing gemmules will have to be secured before a final
decision as to the identity of these several specimens can be
reached.
The spicules of all of them are gently curved and average
around 255 ^ in length. There is probably more variation in
the thickness of the spicules, the range being from 12 ^ to
over 20 p in some of them. The China and Japan specimens
are rather abruptly and bluntly pointed, while the Philippine
specimen is more sharply and gradually pointed.
Flesh spicules. — There are no flesh spicules.
Gemmules. — Annandale found no gemmules in the Yunnan
specimen. He found very few gemmules in the Japanese ma-
terial; and I have not succeeded, after a careful and prolonged
search, in finding even a single gemmule in all of my numerous
specimens of this species representing all three phases from
Lake Biwa. The gemmules are evidently extremely rare.
Those observed by Annandale were at the base of the sponge
attached to the basal membrane, and they very likely are, most
often, left on the support when the sponge is removed. Annan-
dale says concerning the Philippine sponge, "There are very few
gemmules indeed. They occur singly in the basal membrane
and are apparently closely adherent to the support of the sponge.
Each measures about 0.325 mm. in diameter (the shape being
spherical) and is provided with a single straight foraminal tub-
ule on the summit. The granular coat is feebly developed, but
there is a strong outer chitinous coat in continuity with the
basal membrane. The gemmule spicules lie in this coat parallel
to the surface of the gemmule but crossing one another at all
angles."
Gemmule spicules. — "The gemmule spicules are slender, cyl-
indrical, nearly straight. In the middle they bear minute ir-
regular projections, which only take the form of actual spines
towards the two ends. Each end terminates in a stout, straight
spine, surrounded by a row of smaller spines at right angles
to it. None of the spines are retroverted." They are about
one-third of the length of the skeleton spicules.
Type. — The type is in the United States National Museum,
Washington, D. C. I have a small specimen from that mu-
4*1 Gee: Fresh-water Sponges 69
seum labeled as S. dementis, but it differs somewhat from the
typical form of S. dementis as originally described and appears
to be very similar to S. philippinensis.
Distribution. — This species was described from Lake Lanao,
Philippine Islands. Later it was collected in Tali Fu, Yunnan,
China, and then in great abundance and in three distinct phases,
in Lake Biwa, Japan, and in the settling tanks of the waterworks
of the neighboring city of Kyoto. The water supply of Kyoto
comes from Lake Biwa.
Remarks. — Annandale calls attention to the following points
in which S. dementis differs from S. philippinensis. It has
shorter and smoother skeleton spicules; it has a more regular
skeleton; it has a thicker basal membrane; it has adherent gem-
mules with their ill-developed granular coat. I would add,
from Annandale's description, another difference; namely, none
of its gemmule spicules have retroverted spines.
SPONGILLA MICROSCLERIFERA Annandale, 1909. Text fig. 3.
Historical statement. — This sponge was collected by Dr. H.
M. Smith from Lake Taal on the east side of Taal Island, Lu-
zon, December 26, 1907. It was described, without illustra-
tions, by Annandale.9 The United States National Museum has
kindly made available to me a small bit of this sponge, but as
it is so small and there are no gemmules in it I shall quote
parts of the original description, adding my observations where
I have material to justify this; that is, on the skeleton and the
flesh spicules.
Habitat. — The sponge was reported as being abundant around
the shores of the lake and as having been washed up by the
waves on to the shore in large quantities during storms. The
specimens examined by Annandale "appear to have coated both
surfaces of leaves, which have perished and almost disappeared."
General characteristics. — The sponge is "light, fragile, tomen-
tose, . . . apparently without branches and of no great thick-
ness."
Color. — It is "of a dirty white color in dry specimen." In
alcohol the color of my small specimen is a very light brown.
Structure. — "Skeleton practically devoid of spongin but form-
ing a close and almost regular reticulation in which the radiating
and transverse fibers are of approximately equal diameter. The
•Proe. U. S. Nat Mus. 37 (1909) 131-132.
70 The Philippine Journal of Science imi
cS?6£^&^
-^m^^^^^^
FlO. 8. SpongUla tmcrosclerifera Annandale. Skeleton and fresh spicules. No gemmule
spicules were found in my specimens. (Drawing by C. F. Wu.)
free mieroscleres are extraordinarily abundant in the inter-
stices of the skeleton."
Skeleton spicules. — The skeleton spicules are smooth, slightly-
curved, rather slender and of nearly uniform diameter except
near the ends where they become abruptly sharp-pointed.
Scattered among the others in our preparations are a few
heavier spicules which I suppose belong to some other sponge.
Annandale.
Gee.
Length of spicule, fi
254-365
229-310
Diameter of spicule, p.
8.3
6-9
Flesh spicules. — The flesh spicules are very abundant. They
are very variable in length and are extremely thin. As a rule
they are decidedly curved and it is very rare that a straight
one can be found. In the center where the spicule is thickest,
the spines are heaviest and often end in rounded knobs ; toward
the ends of the spicules, where they become extremely tenuous,
the spines are very minute and are sharper pointed. My meas-
urements vary only very slightly from those of Annandale.
Annandale.
Gee.
Length of spicule, p
59.3-124.5
80-127
Diameter in thickest part, fi
1.03-2.07
1.50-3
Gemmules. — "Gemmules few, free, small, spherical, without
a f oraminal tubule, with a thick granular coat, in which the
spicules are arranged tangentially and horizontally in an irreg-
ular manner. Diameter of gemmule 0.35-0.49 mm."
46,1 Gee: Fresh-water Sponges 71
Gemmule spicules. — "Gemmule spicules slender, cylindrical,
nearly straight, bluntly pointed at the ends, irregularly covered
with short, sharp spines, which are more numerous at the ex-
tremities, at which they are usually directed backward, than in
the middle."
Type. — The type is in the United States National Museum
in Washington. I have a small piece of this sponge, without
gemmules, in my collection.
Distribution. — This species has been reported up to date from
only one locality; namely, Taal Lake, Luzon, P. I.
Remarks. — "The most noteworthy characters of this sponge
are the number and hairlike appearance of the free microscleres
which are sometimes of unusual length in spite of their tenuity.
Otherwise there is very little, except perhaps color, to dis-
tinguish it from some forms of Spongilla lacustris. The spec-
imens I have examined are dry and appear to be somewhat
worn on the external surface, but there is no trace of their
having borne branches; the oscula seem to have been fairly
large. The skeleton, in spite of the closeness of its reticulation,
contains much less spongin than is usually the case in Spon-
gilla lacustris, but this is a character liable to a certain amount
of variation, although perhaps less inconstant than is usually
thought."
EPHTDATIA PORTIS Weltner, 1895. Text fiff. 4.
Historical statement. — This sponge was described by Welt-
ner 10 from a specimen collected by Jagor from Libmanan River,
Luzon. Unfortunately there were no drawings to accompany
the original description. Through the kindness of Dr. W.
Arndt, of the University Zoological Museum in Berlin where
the type is preserved, I have been able to secure a very minute
cotype of this sponge. The following description and the il-
lustrations are based upon that material and are supplemented
by a translation from the original description by the author of
the species.
Habitat. — The specimen described by Weltner was found
growing on the leaves of a small water plant, Vallisneria, in
Libmanan River, Luzon.
General characteristics. — My small specimen seems to be a
portion of a very thin crust or film, which was apparently
taken from a plant leaf.
"Archir. fur Naturg. (1895) 141.
72
The Philippine Journal of Science
1931
FIG. 4. Ephydatia fortis Weltner. Spiny skeleton spicules and live of the numerous varia-
tions in the gemmule spicules. (Drawings by C. F. Wu.)
Color. — When dry the sponge is white; the gemmules are also
almost white, but are tinged with a very light brown. In al-
cohol the specimen is nearly transparent.
Structure. — The amount of spongin present is very small and
the sponge appears to be very fragile. The long spicules, singly
or in groups of two or three bound together near their ends,
are woven into large open meshes. A thin basal membrane
still shows the venation of the leaf to which the sponge was
originally attached.
Skeleton spicules. — The large skeleton spicules are spindle-
shaped, thickest in the center, gradually and sharply pointed;
generally slightly curved, a few are straight; nearly always
thickly covered, except near the ends, with minute spines per-
pendicular to the spicule; even the ends are sometimes finely
granular in appearance; only occasionally are smooth or nearly
smooth spicules found and most of these appear to be imma-
ture ones. My measurements show a slightly greater average
length than Weltner's, but this might easily be caused by a
chance selection of the spicules measured, and I consider it a
matter of no special significance.
Weltner.
Gee.
Length of spicules, fi
270-360
297-391
Average, around, /*
30O
325
Diameter of spicules, m
14-16
13-16
Flesh spicules. — There are no flesh spicules in this species.
Gemmules. — The gemmules occur singly in the meshes of the
sponge. They are irregular in shape; some are nearly spherical,
46,1 Gee: Fresh-tvater Sponges 73
while others are somewhat flattened out in one direction making
them appear oblong". They are covered by a layer of birotulate
spicules arranged perpendicularly to the surface of the gemmule,
and in one gemmule the outer rotules were clearly visible as
slight depressions in the thin cuticle that covered the gemmule.
Weltner states that the pore tube is somewhat sunken in the
air-cell layer. He also gives the average diameter of the gem-
mules as 480 /a. I measured two gemmules with the following
results: 340 by 382 ^ and 467 by 510 /x. The first gemmule
measured had its covering spicules disarranged through han-
dling, while in the second the cuticle was not disturbed. The
measurements in both cases included the gemmule spicule layer.
Gemmule spicules. — The gemmule spicules vary a great deal
in length and in general structure. While most of them are
provided with the usual indented terminal rotules characteris-
tic of the genus Ephydatia, yet quite a number of abnormal or
irregular spicules occur. One of the commonest of these is a
long (up to 135 ix or longer), heavily spined spicule that termi-
nates in a large, smooth, sharp spine at each end; at the base
of this spine there is a circle of shaft spines, somewhat larger
than the others, that forms a rudimentary rotule. In some
of the other spicules, which have the normal rotules, the shaft
projects at one or both ends into a sharp point or spine beyond
the rotule. My observations of the normal spicules agree in
detail with those given by Weltner. The spicules vary in length
from about 35 to 65 ^ and the rotules have a diameter of from
20 to 28 /a. Both rotules of a spicule are usually of about the
same diameter. They are irregularly incised, the teeth varying
much in size, and in number from ten to twenty. The shaft is
thickly covered with heavy spines, most of which usually bear
smaller spines. There is much variation in the number of spines
on the shafts of different gemmule spicules, some bear only
eighteen or twenty, while others have as many as forty or more.
Type. — The type of this species is in the University Zoolog-
ical Museum in Berlin. I have a minute cotype in my collection.
Distribution. — This species has been found only in Luzon,
Philippine Islands. The writer has recently described u a va-
riety of this species from specimens collected by Dr. J. R. Baker
from the New Hebrides Islands.
n Ephydatia fortis var. hebridensis, Ann. and Mag. Nat. Hist. X 3
(1929) 28-33, figs.
74 The Philippine Journal of Science
BIBLIOGRAPHY
Weltner, W. Spongillidenstidien III, Katalog und Verbreitung der Be-
kannten Susswasserschwamme. Archiv. fur Naturgesch. 1 (1895).
Annandale, N. Fresh-water sponges in the collection of the United
States National Museum, Part I. Specimens from the Philippines and
Australia. Proc. U. S. Nat. Mus. 36 (1909) 267-632.
Annandale, N. Fresh-water sponges collected in the Philippines by the
Albatross Expedition. Proc. U. S. Nat. Mus. 37 (1909) 131-132.
Annandale, N. Contributions to the fauna of Yunnan. Rec. Indian Mus.
5 (1910) 197. (S. yunnanensis.)
Annandale, N. The sponges of Lake Biwa. Journ. Coll. Sci., Tokyo
Imp. Univ. 39 (1916) 7-11.
Annandale, N. Zoological results of a tour in the Far East. Mem.
Asiatic Soc. Bengal 6 (1918) 201.
ILLUSTRATIONS
TEXT FIGUBES
FIG. 1. Spongilla phUippinensis Annandale. a, Showing both the smooth
and the spined skeleton spicules; also one gemmule spicule (af-
ter Annandale) ; b, smooth skeleton spicules with axial canals
clearly showing. (Drawing by C. F. Wu.)
2. Spongilla dementis Annandale. a, Spined and smooth skeleton
spicules from specimens from Lake Biwa, Japan; the smaller
spicules show the variations in gemmule spicules (after Annan-
dale) ; 6, skeleton and gemmule spicules of specimen from Lake
Lanao, Mindanao; only smooth skeleton spicules were found (af-
ter Annandale).
3. Spongilla micro sclerif era Annandale. Skeleton and fresh spicules.
No gemmule spicules were found in my specimens. (Drawing
by C. F. Wu.) *
4. Ephydatia fortis Weltner. Spiny skeleton spicules and five of the
numerous variations in the gemmule spicules. (Drawings by C.
F. Wu.)
75
PLANKTON DIATOMS PROM VLADIVOSTOK BAY
By B. W. Skvortzow
Of Harbin, China
TWO PLATES
The diatoms included in this paper were collected by Mr. N.
E. Kabanov and me in Vladivostok Bay July 18, 1928. So far
as I know, there is no published list of plankton diatoms from
this part of the Sea of Japan and the enumeration will be of
interest. The number of forms in the present collection is not
great, but there are some interesting ones. I am describing a
species of Synedra as new.
LEPTOCYLINDRUS DANICUS Clcve. Plate 2, fi*. 3.
Clbve, Pelag. Diat. fr. Kattegat (1889) 54; Planktonundersokningar
Cilioflag. och Diat. (1894) 15, pis. 1, 2, figs. 4, 5; Hustedt in A.
Schmidt, Atlas Diatom. (1920) pi. 321, fig. 12; Kieselalgen (1929)
558-59, figs. 318, 319.
Cells cylindrical, with flat ends, forming filaments. Valves
without structure. Chromatophores numerous. Diameter of
filaments, 0.007 to 0.008 millimeter. Geographic distribution:
Atlantic and Pacific Oceans.
SCELETONEMA COSTATUM (Grev.) Cleve. Plate 2, fi*. 4.
Cleve, Bih. Kongl. Sv. Vet.-Akad. Handl. 5 (1878) 18; A. Schmidt,
Atlas Diatom. (1892) pi. 180, figs. 41-45; (1920) pi. 321, figs. 5, 6;
Husteot, Kieselalgen (1928) 311-13, fig. 149.
Cells 0.007 to 0.009 millimeter broad, 0.018 to 0.022 in length.
Geographic distribution: A typical pelagic diatom known from
the Atlantic and Pacific Oceans.
DITTLIUM BRIGHTWELII (West) Gran.
V. Heurck, Synopsis (1885) pi. 114, figs. 3-9.
Cells 0.12 to 0.15 millimeter in length, 0.025 to 0.035 in
breadth. Geographic distribution: Common in plankton of
oceans, known from the Sea of Japan.
77
78 The Philippine Journal of Science 1931
CHAETOCERAS SOCIALE Lander. Plate 1, fig. 2.
Lauder, Diatom. Hong Kong (1864) 77, pi. 8, fig. 1.
I am giving here the original diagnosis of this species, from
Henry Scott Lauder :
Filaments slender, aggregated, embedded in gelatine, with wiry spirally
dotted awns, some of which are more elongate and converge to a common
centre. This is the smallest species I have seen. By the aggregation of
the filaments in gelatine, it forms roundish, flattened fronds. Frustules
quadrate with an awn from a little within each angle, one of them being
more elongated, varying in the length, according to the distance of the
frustules, to a common centre, to which the elongated awns converge:
many frustules, however, occur in which the awns are not thus connected:
side view oval.
Our specimens were 0.008 to 0.01 millimeter in diameter.
Geographic distribution: Atlantic and Pacific Oceans; known
from Japanese waters.
CHAETOCERAS CRIOPHILUM Castracane forma VOLANS (Schatt) Gran. Plate 1, fisrs.
5 and 6.
Castracane, Diatom. Challenger (1886) 78; Gran, Diatom. Arct.
Meere (1904) 532-33, fig. 4; Nord. Plankton (1906) 71, fig. 85;
Gran and Yendo, Japan Diatom. (1914) 7; Okamura, Littor. Dia-
tom. Japan (1911) 90, pi. 3, figs. 33-37; Karsten, Phytopl. Antarkt.
Meeres (1905) 118, pi. 15, fig. 8; PeragauU), Diatom. Mar. France
(1908) 475; Hustedt in A. Schmidt, Atlas Diatom. (1920) pi. 342,
figs. 2, 3.
Cells solitary, 0.02 to 0.023 millimeter broad, from front view
quadrangular with angles. Setse very robust, curved, covered
with solid spines. Geographic distribution: Atlantic and Pa-
cific Oceans; the Sea of Japan.
CHAETOCERAS COMPRESSUM Lander. Plate 1, fi*. 1.
Lauder, Diatom. Hong Kong (1864) 78, pi. 8, fig. 6.
Chain straight, 0.01 to 0.018 millimeter broad. Spores with
verrucose dots on the margin. Geographic distribution: Com-
mon in Atlantic and Pacific Oceans. Reported from the Sea of
Japan.
CHAETOCERAS COMPRESSUM var. GRACILIS Hustedt. Plate 1, fig:. 7.
Hustedt, in A. Schmidt, Atlas Diatom. (1921) pi. 338, fig. 7.
Chain thinner, 0.006 to 0.008 millimeter broad. Geographic
distribution: Known only from the Sea of Japan.
CHAETOCERAS DIDYMUM Ehrenb. var. ANGLICA Gran. Plate 1, fig. 3.
Gran, Nord. Plankton (1906) 80, fig. 95.
Cells forming a straight chain, 0.02 to 0.03 millimeter broad.
Cells in the front view rectangular with a large dot in the middle
46,1 Skvortzow: Diatoms from Vladivostock Bay 79
part. Setae start at a little inside of the margin. Geographic
distribution: Atlantic and Pacific Oceans; Sea of Japan.
CHAETOCERAS GRACILE Schtttt. Plate 1, fi*. 10.
Schutt, Chaetocera and Peragallo. (1895) 42, figs. IZa-d.
Chaetoceras septentrionale Oestrup, Mar. Diatom. Gronland. (1895)
457, pi. 7, fig. 88; Gran, Diatom. Arkt. Meere (1904) 542; Paulsen,
On some Perid. a. Plankt. Diatom. (1905) 5-6, fig. 6.
Cells solitary, 0.015 to 0.025 millimeter broad; in the front
quadrangular and in valve view elliptical. Setae thin, very long.
Geographic distribution: Atlantic Ocean.
CHAETOCERAS DECIPIENS Clere. Plate 1, fig. 8.
Clevb, Diatom. Arct. Sea (1873) 11, fig. 5.
Chain straight, with rectangular cells, 0.035 to 0.55 millime-
ter broad. Geographic distribution : Atlantic and Pacific Oceans ;
Sea of Japan.
CHAETOCERAS CONSTRICTUM Gran. Plate 2, fig. 2.
Gran, Tret. Phytopl. N. Atlant. (1897) 17, pis. 11-13, pi. 3, fig. 42;
Nord. Plankton (1906) 80, fig. 96; Okamura, Chaetoceras and Pe-
rag. Japan (1907) 96, pi. 4, figs. 64a, b; Peragallo, Diatom. Mar.
France (1908) 491, pi. 134, fig. 5; Hustedt in A. Schmidt, Atlas
Diatom. (1921) pi. 338, fig. 1.
Chain straight, 0.01 to 0.012 millimeter broad. Cells in a
front view rectangular with slightly projecting angles, valves
concave, foramina lanceolate, constricted. Girdle band one-third
of the cell height. Setae thin, starting from the angles of the
valve, crossing one another close to their insertions, diverging
at an obtuse angle. Terminal setae are not differentiated.
Geographic distribution: Atlantic and Pacific Oceans.
Okamura reports this species from the Kuriles.
CHAETOCERAS LACINIOSUM Schutt. Plate 2, fig. 1.
Schutt, Arten Chaetoceras u. Peragallo. (1895) 38, figs. 5a-c; Gran,
Phytopl. N. Atlant. (1897) 17, figs. 4-7; Nord. Plankton, (1906)
82, fig. 99; GRAN and Yendo, Japan Diatom. (1914) 18-19, fig. 11.
Chaetoceras distatis Clevb, Plankton. Cilioll. och. Diatom. (1894)
14, pi. 2, fig. 3.
Chaetoceras distans Ostenfeld, Flora Koh-Chang (1902) 255, fig. 13.
Chaetoceras commutatum Cleve, Plankton. Vegetabil. (1896) 28, figs.
9, 10.
Chaetoceras ostenfeldii Cleve, Plankt. N. Sea (1900) 21, pi. 8, fig. 19.
Chaetoceras pelagicum Cleve, Diatom Arct. Sea (1873) 11, pi. 1, fig.
4.
80 The Philippine Journal of Science 1931
Chaetoceras distans var. laciniosa Schutt in Peragallo, Diatom. Mar.
France (1908) 483, pi. 132, fig. 6; Ikari, Chaetoceras Japan (1928)
253-54, fig. 8a.
Chain straight, 0.01 to 0.015 millimeter broad, composed of
many cells. Foramina large, as long as the cell height and ellip-
tical, slightly constricted in the middle. Valves in a front view
rectangular with projecting angles. Girdle band rather longer,
about two-thirds of the cell height. Setae thin, starting from
the angles of the valve, crossing one another close to their in-
sertions, diverging at an obtuse angle. Terminal setae are dis-
posed parallel. Geographic distribution: Common in Atlantic
and Pacific Oceans. Known in Japanese waters from Oshoro,
Takashima, Ajiro, Naha, Volcano Bay, Misume, and Seto.
CHAETOCERAS sp. Plate 1, fiff. 4.
Chain straight, 0.012 to 0.015 millimeter wide. Cells in a
front view rectangular, valves concave, foramina lanceolate.
Girdle band rather longer, about one-third of the cell height.
Setae thin, irregularly disposed, terminal setae long, more or less
divergent.
THALASSIOTHRIX NITZSCHIOIDES Gran. Plate 2, figs. 7 and 8.
V. Heurck, Synopsis (1883) 43, fig. 7.
A common pelagic diatom forming zigzag clusters. Cells lan-
ceolate, 0.012 to 0.074 millimeter in length, 0.0025 to 0.003 in
breadth with marginal striae 12 to 15 in 0.01 millimeter. Geo-
graphic distribution: North Atlantic and Pacific Oceans.
THALASSIOTHRIX FRAUENFELDII Gran. Plate 2, fiff. 9.
V. Heurck, Synopsis (1883) pi. 37, figs. 11-13.
Cells forming star-shaped or zigzag clusters. Cells 0.002 to
0.15 millimeter in length. Geographic distribution : Atlantic and
Pacific Oceans. Sea of Japan.
THALASSIOTHRIX ANTARCTICA Schimper. Plate 2, figs. 5 and 6.
Karsten, Phytopl. Atlant. Meeres (1905) 124, pi. 17, fig. 12.
Cell filiform, curved, 1.32 to 1.65 millimeters in length, 0.0037
to 0.005 in breadth. Striae 14 to 16 in 0.01 millimeter. Geo-
graphic distribution: Atlantic and Pacific Oceans.
ASTERIONELLA JAPONICA Cleve. Plate 2, figs. 10 and 11.
Okamura, Littor. Diatom. Japan (1911) 11, pi. 13, fig. 56.
A typical pelagic diatom forming star-shaped and zigzag
clusters. Cells length 0.045 to 0.144 millimeter. Geographic
distribution: Atlantic and Pacific Oceans; Sea of Japan.
46, i Skvortzow: Diatoms from Vladivostock Bay 81
SYNEDRA AFFINIS Kutz. var. GRACILIS Grun.
V. Hburck, Synopsis (1883) pi. 41,. fig. 15B; Peragallo, Diatom. Mar.
France (1908) 320, pi. 130, figs. 23, 24.
Valve lanceolate, in the middle part inflated. Length, 0.35 to
0.44 millimeter; breadth, 0.012; striae 12 in 0.01 millimeter.
Geographic distribution : A littoral diatom known from the At-
lantic and Pacific Oceans and Mediterranean Sea.
SYNEDRA JAPONICA sp. nov. Plate 2, fig. 13.
Cell free, linear or linear-lanceolate, pseudoraphe distinct.
Valve 0.48 to 0.59 millimeter in length, in the middle part,
0.0034 to 0.0045 in breadth; striae 15 in 0.01 millimeter.
RHIZOSOLENIA ALATA Bright, forma GRACILLIMA (Cleve) Grun.
Peragallo, Monogr. Rhizosolenia (1892) 20, pi. 5, fig. 12; Hustedt
in A. Schmidt, Atlas Diatom. (1920) pi. 317, figs. 8-10.
Cells 0.85 to 0.88 millimeter in length and 0.0074 in breadth.
Geographic distribution: Common in plankton in Atlantic and
Pacific Oceans.
RHIZOSOLENIA SETIGERA Bright.
Peragallo, Monogr. Rhizosolenia (1892) 17, pi. 4, fig. 12-16.
Cell linear, hyaline, 0.65 to 0.75 millimeter in length and
0.011 in breadth. Spine thin, 0.04 to 0.07 millimeter in length.
Geographic distribution: Atlantic, Pacific and Indian Oceans,
Sea of Japan.
NITZCHIELLA LONGISSIMA (Breb.) Ralfs forma PARVA V. Heurck. Plate 2, figr. 12.
V. Heurck, Synopsis (1883) pi. 70, fig. 3.
Length, 0.118 to 0.2 millimeter; breadth in the middle part,
0.0035 to 0.004. Geographic distribution : Littoral diatom, cos-
mopolitan.
PLEUROSIGMA FASCIOLA Ehrenb. var. ARCUATUM Donk. Plate 1, fiff. 9.
Pleurosigma arcuatum Donkin, T. M. S. (1858) 25 fig. 10; Cleve,
Synopsis Navic. Diatoms (1894) Part I, II;, Peragallo, Monogr.
Pleurosigma (1890 to 1891) 26, pi. 8, figs. 34, 35.
Valve lanceolate, with produced beak-shaped ends. Length,
0.1 to 0.15 millimeter; breadth, 0.015 to 0.018. Geographic dis-
tribution: A littoral diatom known from the Atlantic and Pa-
cific Oceans.
259737 6
ILLUSTRATIONS
Plate 1
Fig. 1. Chaetoceras compression Lauder.
2. Chaetoceras sociale Lauder.
3. Chaetoceras didymum Ehrenb. var. anglica Gran.
4. Chaetoceras sp.
5. Chaetoceras criophilum Castr. f. volans (Schiitt) Gran.
6. Chaetoceras criophilum Castr. f. volans (Schiitt) Gran.
7. Chaetoceras compressum Lauder var. gracilis Hustedt.
8. Chaetoceras decipiens Cleve.
9. Pleurosigma fasciola Ehrenb. var. arcuatwn Donk.
10. Chaetoceras gracile Schiitt.
Plate 2
Fig. 1. Chaetoceras laciniosum Schiitt.
2. Chaetoceras constrictum Gran.
3. Leptocylindrus danicus Cleve.
4. Sceletonema costatum (Grev.) Cleve.
5. Thalassiothrix antarctica Schimper.
6. Thalassiothrix antarctica Schimper.
7. Thalassiothrix nitzschioides Grun.
8. Thalassiothrix nitzschioides Grun.
9. Thalassiothrix frauenfeldii Grun.
10. Asterionella japonica Cleve.
11. Asterionella japonica Cleve.
12. Nitzchiella longissima (Breb.) Ralfs f. parva V. Heurck.
13. Synedra japonica sp. nov.
88
Skvortzow: Diatoms from Vladivostok Bay.]
[Philip. Journ. Scl, 46, No. 1.
PLATE 1.
Skvortzow : Diatoms from Vladivostok Bay.]
[Philip. Journ. Sci., 46, No. 1.
6 5
PLATE 2.
MYCETOZOA FROM NORTH MANCHURIA, CHINA 1
By B. W. Skvortzow
Of Harbin, China
FIVE PLATES
A certain amount of work has been done on the Mycetozoa of
some parts of Europe and the western parts of Asia, but the
slime fungi of eastern Asia, especially of Manchuria, have been
completely overlooked. Contributions to the Mycetozoa of Si-
beria have been published by Dr. N. N. Lavrov, of the Tomsk
University of Siberia, USSR, and the Japanese Mycetozoa were
studied by A. and G. Lister, in Mycetozoa from Japan ;2 and by
K. Ninakata, in the list of Japanese Myxomycetes.3
The Mycetozoa that form the subject of this paper were col-
lected by the author in North Manchuria in 1920 to 1929, es-
pecially in Harbin and in the mountainous part of the country
near Erth'enkiangtzu and Maoershan station on the Chinese
Eastern Railway. The number of forms found in the present
collection is not great, but there are some interesting ones. In
this note thirty-two slime fungi are enumerated and ten are
described as new; namely, Badhamia mandshurica, Physarwm
compressum, Ph. griseum, Ph. mandshuricum, Ph. asiaticum,
Diderma rugosum Macb. var. asiatica, Lepidoderma mandshurica,
Licea brassica, L. mandshurica, and Trichia asiatica.
The Mycetozoa are cosmopolitan in their distribution and the
finding of new species in the eastern part of Asia is of scienti-
fic interest. Most Manchurian forms of Mycetozoa are charac-
teristic of temperate regions; such are, Fuligo septica, F.
muscorum, Diderma spumarioides, Stemonitis splendens, S.
herbatica, Lycogala epidendrum, Trichia persimilis, T. contorta,
Hemitrichia clavata, Arcyria denudata, A. cinerea, and Pericha-
ena depressa. Some species of slime fungi, Ceratiomyxa fruti-
1 From the laboratory of natural history of the Third High School of the
Chinese Eastern Railway Co., Harbin, China. (Formerly the Commercial
School.)
'Journ. of Bot. 42 (1904) 97-99, t. 458; 42 (1906) 227-230.
8Bot. Mag. Tokyo 42 (1908) 317.
85
86 Tfee Philippine Journal of Science 1931
culosa, Didymium dubium, Licea biforis, Arcyria earnea, and
others, are here recorded from Asia for the first time. All
Physarum species found in Manchuria are identified as new to
science.
This note is illustrated by diagrams by the author and by
photographs made by lea mikro-camera with Planar 1 : 4.5
F = 2 cm of Carl Zeiss.
CERATIOMYXA FRUTICULOSA Macbr. var. FLEXUOSA Lister. Plate 4, fig. 3.
Sporophores long, branching, white, 1 to 3 millimeters high.
Branches of sporophores 30 to 60 microns, thickly covered with
spines. Spores white, smooth, ovoid, 8 to 13 microns long.
Habit : On rotten wood ; near Maoershan station, Chinese Eastern
Railway, August, 1928. Geographic distribution: Abundant in
the Tropics ; recorded in Europe, Japan, and South Africa.
BADHAMIA MANDSHURICA sp. nov. Plate 2, figs. 6, 7, and 8.
Plasmodium unknown. Sporangia subglobose, sessile, minute,
0.2 to 0.5 millimeter in diameter, scattered or in small clusters,
black, somewhat rugose and gray; sporangium wall membra-
nous, with scanty deposits of lime granules. Capillitium a net-
work of slender threads with white lime deposits. Spores free,
blackish brown, round, somewhat minutely spinulose, 13 to 15.5
microns in length. Habit: On the bark of trees; Harbin, Au-
gust, 1920.
PHYSARUM COMPRESSUM sp. nov. Plate 1, figs. 7, 8, and 9.
Sporangia stalked, gregarious, discoid or compressed, some-
times umbilicate above, 0.4 to 1 millimeter in diameter, grayish
white, rugulose; sporangium wall membranous, with abundant
deposits of white lime granules. Stalk furrowed, yellow-brown.
Capillitium a persistent network of stout, rigid, hyaline threads
and numerous rounded dark yellow-brown lime knots. Spores
11 to 12 microns in diameter, brown, spinulose. Habit: On dead
wood; Harbin, August, 1920.
PHYSARUM GRISEUM sp. nov. Plate 1, figs. 4, 5, and 6.
Sporangia sessile, subglobose or elongate, clustered, grayish
white, 0.4 to 0.7 millimeter in diameter, rugulose; sporangium
wall membranous, with lime granules. Capillitium consisting
of short hyaline threads connected by angular branching brown-
yellow lime knots. Spores purplish brown, spinulose, 9.2 to
12 microns in diameter. Habit: On dead wood; Harbin, No-
vember, 1920.
46,1 Skvortzow: Mycetozoa from Manchuria 87
PHYSARUM MANDSHURICUM sp. nov. Plate 1, figs. 1, 2, and 3.
Sporangia subglobose, reniform, stalked, erect or somewhat
inclined, scattered or clustered, two or more often borne on a
single stalk, 0.3 to 0.7 millimeter in diameter, white, rugose;
sporangium wall membranous, with white granules. Stalk sub-
ulate or cylindrical, furrowed, 1 millimeter long, yellow-brown,
usually free from refuse matter. Capillitium a network of color-
less branching threads, lime knots large, not numerous. Spores
dark reddish brown, spinulose, 11 to 12 microns in diameter.
Habit : On dead bark of trees ; Maoershan station, Chinese East-
ern Railway, August, 1928.
PHYSARUM ASIATICUM sp. nov. Plate 2, figs. 9, 10, and 11.
Sporangia subglobose or irregularly ovoid, 0.2 to 0.5 milli-
meter in diameter, sessile, heaped or gregarious, rugose, whitish
black; sporangium wall membranous, with dense included clus-
ters of minute white lime granules. Capillitium a network of
dark brown threads, with irregular dark brown lime knots.
Spores dark violet-brown, 10 to 12 microns in diameter, spinu-
lose. Habit: On bark of trees; Harbin, September, 1920.
FULIGO SEPTICA Graelin. Plate 3, fig. 3; Plate 4, fig. 4.
iEthalia pulvinate, 0.5 to 7 centimeters broad, light yellow.
Capillitium consisting of a loose network with yellow lime
knots. Spores violet, smooth, 6.8 to 9.5 microns in diameter.
Habit: On dead wood and earth; Harbin, July, August, and
September, 1929. Geographic distribution: Abundant in tem-
perate and tropical regions.
FULIGO SEPTICA Gmelin var. RUPA R. E. Pries. Plate 3, fig. 2.
.ZEthalia pulvinate, 2 to 4 centimeters in diameter, brick red
or yellow-red. Capillitium scanty, consisting of a loose network
of slender hyaline threads. Spores violet-brown, 6.8 to 9.6 mi-
crons in diameter. Habit: On earth and on leaves; near Maoer-
shan station, Chinese Eastern Railway, August, 1928. Geogra-
phic distribution: Abundant in temperate and tropical regions.
FULIGO MUSCORUM Alb. and Schwein.
JEthalia pulvinate, 0.4 to 0.8 millimeter in diameter, scattered,
yellow. Capillitium of numerous irregular large orange lime
knots. Spores violet-brown, spinulose, 10.2 to 12 microns in
diameter. Habit: On bark of Cladrastis amurensis; Maoershan
station, Chinese Eastern Railway, August, 1928. Geographic
distribution : Europe and Ceylon.
88 The Philippine Journal of Science mi
DIDERMA GLOBOSUM Pers. Plate 4, fi«r. 2.
Sporangia subglobose, sessile, forming large colonies, 0.2 to
0.5 millimeter in diameter, white-gray; sporangium wall of two
layers, the outer eggshell-like, composed of globular lime gran-
ules. Columella indistinct. Capillitium dark brown branched
threads. Spores dark black-brown, fine spinulose, 11 to 11.5
microns in diameter. Habit: On leaves and twigs of Car ex sp.
and Artemisia sp.; near Maoershan station, Chinese Eastern
Kailway, August, 1928. Geographic distribution : Recorded from
Europe, United States, and British Columbia.
DIDERMA SPUMARIOIDES Fries.
Sporangia crowded, forming colonies, globose, sessile, 0.5 to
1.5 millimeters in diameter, smooth, white. Sporangium wall of
two layers. Capillitium slender threads. Spores dark reddish
brown, spinulose, 7.4 to 11.1 microns in diameter. Habit: On
dead wood; Harbin, August, 1920. Geographic distribution:
Europe, United States, Canada, Ceylon, Japan, West Indies,
Bermuda, and southern Chile.
DIDERMA RUGOSUM Macbride var. ASIATICA var. nov.
Plasmodium gray. Sporangia stalked, subglobose, 0.4 to 0.5
millimeter in diameter, grayish white, reticulated, wrinkled.
Sporangium wall single, with deposits of lime in minute granules.
Stalk 0.5 to 0.7 millimeter high, furrowed, yellow-brown. Colu-
mella clavate, about half the height of the sporangium. Capil-
litium consisting of slender colorless threads, anastomosing and
branching towards the tips. Spores purplish brown, minutely
warted, 7.5 to 9 microns in diameter. Habit: On leaves of
Brassica chinensis; Harbin, August, 1928. Geographic distrib-
ution : The typical Diderma rugoswm was recorded from Europe,
Ceylon, Japan, America, and United States.
LEPIDODERMA MANDSHURICA sp. nov. Plate 2, figs. 1, 2, and 3; Plate 5, fig. 2.
Sporangia forming short, subglobose or elongate pulvinate
plasmodiocarps, 0.5 millimeter to 5 centimeters long, 0.5 to 5
millimeters broad, silvery gray, clothed with brilliant crystalline
scales of lime ; capillitium of slender brownish threads, branched
and anastomosing. Spores brown-violet, smooth, 6.8 to 7.2 mi-
crons in diameter. Habit : On leaves and dead twigs, Maoershan
station, Chinese Eastern Railway, August, 1928.
STEMONITIS SPLENDENS Rost. var. FLACCIDA Lister.
Sporangia cylindrical, obtuse, stalked, dark brown, adhering
to each other, forming large colonies. Stalk black. Total height
46,i Skvortzow: Mycetozoa from Manchuria 89
1 to 1.8 centimeters. Capillitium of purplish brown branching
threads. Spores dark brown, 7.4 to 7.8 microns in diameter.
Habit: On dead wood; Erth'enkiangtzu station, Chinese Eastern
Railway, July, 1927. Geographic distribution: Europe and
America.
STEMONITIS HERBATICA Peck.
Sporangia cylindrical, closely clustered, 8 to 10 millimeters
high, brown. Stalks 2 to 3.5 millimeters high. Capillitium
brown threads, forming a loose network. Spores smooth, 5.1
to 5.4 microns in diameter. Habit: On dead wood; Maoershan
station, Chinese Eastern Railway, August, 1928. Geographic
distribution: Europe, United States, Ceylon, and Japan.
DICTYDIUM CANCELLATUM Macbr.
Sporangia subglobose, dark red-brown, stalked, formed of
numerous ribs, connected by slender transverse threads, 1.5 to 2
millimeters high and 0.3 to 0.5 millimeter broad. Spores red,
5.2 to 5.7 microns in diameter, minutely warted. Habit: On
dead wood; Harbin, August, 1920. Geographic distribution:
Europe, Africa, America, and Japan.
LICEA BIFORIS Morgan.
Sporangia scattered, ellipsoid, elongate, sessile, 0.1 to 0.3 milli-
meter long, 0.06 to 0.1 millimeter broad, yellow-brown. Spores
round, smooth, light yellow, 9.2 to 11.1 microns in diameter,
with oil drops. Habit: On dead bark; Harbin, November, 1929.
Geographic distribution: Japan, Pennsylvania, Ohio, and Can-
ada.
LICEA BRASSICA sp. nov. Plate S, fig. 4.
Sporangia scattered, depressed, forming straight, curved, or
branching plasmodiocarps 0.5 to 5 millimeters long, grayish
white, reticulated and little wrinkled. Sporangium wall single,
with deposits of lime in minute granules. Spores purplish
brown, smooth, 9.2 to 9.5 microns in diameter. Habit: On
leaves of Brassica chinensis. This species somewhat resem-
bles Licea flexuosa Pers., but differs from it in the color of the
sporangium wall and by the spores.
LICEA MANDSHURICA sp. nov. Plate 2, figs. 4 and 5.
Sporangia sessile, depressed, forming straight, curved, and
wrinkled plasmodiocarps, 1.5 millimeters long, olive or dark
gray, more or less closely covered with flat, rounded, angular
crystalline scales of lime. Capillitium and columella wanting.
90 The Philippine Journal of Science imi
Spores dark violet-brown, 6 to 7.2 microns in diameter, nearly-
smooth. Habit: On earth; Harbin, August, 1929.
LYCOGALA EPIDENDRUM Fries.
iEthalia subglobose, 5 to 6 millimeters in diameter, dark
gray, warted. Pseudocapillitium in the form of tubes, marked
with close transverse wrinkles. Spores gray, spinulose, 6.8
to 8.5 microns in diameter. Habit: On dead wood; Harbin,
August, 1920. Geographic distribution: British Isles and fre-
quent in all temperate and tropical regions.
TRICHIA PERSIMILIS Karst.
Sporangia globose, sessile, forming large colonies, 0.3 to 0.6
millimeter in diameter, yellow-brown. Capillitium 3.8 to 4
microns in diameter, with spiral bands and short spines.
Spores yellow, 11.1 to 11.4 microns in diameter, with pitted
warts. Habit: On rotten wood; Maoershan station, Chinese
Eastern Railway, August, 1928. Geographic distribution : Brit-
ish Isles, Europe, Ceylon, Java, and Peru.
TRICHIA VARIA Pers.
Sporangia globose, ovoid, only sessile, 0.5 to 1 millimeter
in diameter, forming large colonies. Sporangium wall mem-
branous, pale yellow. Capillitium yellow, elater 3.5 to 4.5 mi-
crons in diameter, tapering at the ends, with two spiral bands.
Spores yellow, minutely warted, 11 to 13 microns in diameter.
Habit: On dead wood; Harbin, November, 1920. Geographic
distribution : Europe, United States, Ceylon, and northern India.
TRICHIA CONTORTA Rost. var. INCONSPICUA Lister.
Sporangia clustered, forming large colonies; 0.2 to 0.7 milli-
meter in diameter, sessile, brown ; sporangium wall membranous,
reddish brown. Capillitium simple, elaters with four distinct
spiral bands, 3.7 to 4 microns in diameter. Spores yellow,
minutely spinulose, 11 to 13 microns in diameter. Habit: On
bark of trees; Harbin, 1920. Geographic distribution: Widely
distributed throughout north temperate regions.
TRICHIA ASIATICA sp. nov. Plate 1, fiffg. 10, 11, and 12.
Sporangia globose, usually crowded and seated on a common
membranous hypothallus, 0.5 to 0.8 millimeter in diameter,
brown or yellow-brown, shining. Capillitium and spores in
mass yellow; sporangium wall membranous, yellow. Capilli-
tium of bright yellow or orange elaters, 3.5 to 4 microns in
diameter, with four bands, forming a close spiral studded with
46, i Skvortzoiv: Mycetozoa from Manchuria 91
many spines. Spores dark yellow-brown, minutely warted, 10
to 12.5 microns in diameter. Habit: On bark of trees; Harbin,
November, 1929.
HEMITRICHIA SERPULA Rosr. Plate 5, fig. 1.
Sporangia forming winding branched plasmodiocarps, 0.2 to
0.5 millimeter wide, uniting into a close net, golden yellow;
sporangium wall membranous, yellow. Capillitium elastic yel-
low threads, 4 to 4.5 microns in diameter, marked with spiral
bands, spinose. Spores yellow, reticulated with bands, forming
a regular net, 11 to 11.5 microns in diameter. Habit: On bark
of trees ; Maoershan station, Chinese Eastern Railway, August,
1928. Geographic distribution: Europe, abundant in Japan,'
United States, the Tropics, Australia, New Zealand, and the
Cape Province.
HEMITRICHIA VESPARIUM Macbr.
Sporangia clavate, stalked, crowded, 1 to 1.4 millimeters high,
dark red. Stalks combined in clusters of from five to ten,
red. Capillitium red, twisting threads, 5.7 to 6 microns in
diameter, with spiral bands and numerous scattered spines.
Spores red, warted, 11 microns in diameter. Habit: On dead
wood; Harbin, October, 1920. Geographic distribution: Re-
corded from most temperate and tropical regions ; United States.
HEMITRICHIA CLAVATA Rost.
Sporangia stalked, gregarious or crowded, 0.9 to 2 milli-
meters high, olivaceous-yellow; sporangium wall yellow. Stalk
red-brown. Capillitium a network of branched yellowish
threads, 5.3 to 5.7 microns in diameter, marked with five spiral
bands, without spines. Capillitium ends subclavate. Spores
dark brown, warted, 7.2 to 7.6 microns in diameter. Habit:
On dead wood; Maoershan station, Chinese Eastern Railway,
August, 1928. Geographic distribution : Widely distributed and
common in all temperate and tropical regions.
ARCYRIA DENUDATA Wettstein.
Sporangia stalked, ovoid or cylindrical, 0.6 to 2 millimeters
high, 0.4 to 1.2 millimeters broad, reddish or reddish brown,
Capillitium a close elastic network of red threads with thicken-
ings of cogs or spines and half-rings. Spores red, smooth, 6
to 7 microns in diameter. Habit: On dead wood and bark of
trees; Maoershan station, Chinese Eastern Railway, August,
1928. Geographic distribution: Abundant in temperate and
tropical regions.
92 The Philippine Journal of Science
ARCYRIA CINEREA Pers. Plate 4, fiff. 1.
Sporangia stalked, almost gregarious, cylindrical, 0.7 to 1.2
millimeters long, 0.4 to 0.7 millimeter in diameter; pale gray.
Stalk 0.4 to 0.6 millimeter long, dark gray. Capillitium a
close network of gray threads, warted and spinulose. Spores
gray, smooth, 6 to 6.5 microns in diameter. Habit: On dead
wood ; Harbin, November, 1929. Geographic distribution : Com-
mon and widely distributed in temperate regions.
ARCYRIA CARNEA G. Lister.
Sporangia stalked, clustered, ovoid or shortly cylindrical,
flesh-colored, 1.5 to 2 microns high; cup plaited at the base,
red. Stalks black. Capillitium a close and only slightly elas-
tic network of dark red threads, 3.7 to 4.5 microns in diameter,
marked with a loose spiral of flat-tipped cogs or spines. Spores
reddish, 6.8 to 7.4 microns in diameter, smooth. Habit: On
dead wood; Maoershan station, Chinese Eastern Railway,
August, 1928. Geographic distribution: Europe and Japan.
PERICHAENA DEPRESSA Libert.
Sporangia sessile, crowded, flattened, 0.1 to 0.5 millimeter
in diameter, sometimes forming short branching plasmodio-
carps of 1 to 3 millimeters diameter, purple-brown. Sporan-
gium wall of two layers. Capillitium branched yellow threads,
minutely warted. Spores yellow, warted, 9 to 11 microns in
diameter. Habit: On dead bark of Populus simonii; Harbin,
November, 1929. Geographic distribution: Widely distributed
in temperate and tropical regions.
ILLUSTRATIONS
Plate 1
Figs. 1 to 3. Physarum mandshuricum sp. no v.; 1, sporangia, X 20; 2
and 8, capillitium and spore, X 500.
4 to 6. Physarum griseum sp. nov.; 4, sporangia, X 20; 5 and 6,
capillitium and spore, X 500.
7 to 9. Physarum compressum sp. nov.; 7, sporangia, X 20; 8 and
9, capillitium and spore, X 500.
10 to 12. Trichia asiatica sp. nov.; 10, sporangia, X 20; 11 and 12,
capillitium and spore, X 500.
Plate 2
Figs. 1 to 3. Lepidoderma mandshurica sp. nov.; 1, sporangia, X 20; 2
and 8, capillitium and spore, X 500.
4 and 5. Licea mandshurica sp. nov.; 4, sporangia, X 20; 5, spores,
X 500.
6 to 8. Badhamia mandshurica sp. nov.; 6, sporangia, X 20; 7 and
8, capillitium and spore, X 500.
9 to 11. Physarum asiaticum sp. nov.; 9, sporangia, X 20; 10 and 11,
capillitium and spore, X 500.
Plate 3
Fig. 1. Trichia persimiLis Karst., sporangia, X 15.
2. Fuligo septica Gmelin var. rufa R. E. Fries, aethalia, X 1. 5.
3. Fuligo septica Gmelin, aethalia, X 1- 5.
4. Licea brassica sp. nov., sporangia, X 10.
Plate 4
Fig. 1. Arcyria cinerea Pers., sporangia, X 15.
2. Diderma globosum Pers., sporangia, X 15.
3. Ceratiomyxa fruticulosa Macbr. var. flexuosa Lister, sporophores,
X15.
4. Fuligo septica Gmelin, aethalia, X 1. 5.
Plate 5
Fig. 1. Hemitrichia serpula Host., sporangia, X 8.
2. Lepidoderma mandshurica sp. nov., sporangia, X 5.
93
Skvoutzow: Myoktozoa from Manchuria. J
[Philip. Journ. So.. 48, No. 1.
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PLATE 5.
PELAGIC DIATOMS OF KOREAN STRAIT OF THE
SEA OF JAPAN
By B. W. Skvortzow
Of Harbin, China
TEN PLATES
Our knowledge of pelagic diatoms of the Pacific Ocean is still
incomplete in comparison with that of the northern part of the
Atlantic. A certain amount of work has been done on the
marine diatoms of various parts of Japan, but most of the algse
of the Korean shores have been overlooked. Contributions on
the diatoms of the Sea of Japan have been published by K. Oka-
mura, J. Ikari, H. H. Gran, K. Yendo, B. Schroder, and F.
Hustedt.
The diatoms that form the subject of this memoir were ob-
tained from material collected by the Government Fishery Ex-
perimental Station, of Fusan, Chosen, and sent me through the
kindness of Dr. Yojiro Wakiya, director of that station. The
gatherings were the following:
1. Fusan, between Katokuto Island and Tatapo, January 12, 1925.
2. Port Fusan, between Fusan and Zetsueito Island, November 9, 1925.
3. Near Port Fusan, February 28, 1926.
4. Northern Tsushima Island, September 13, 1925.
5. Near Tsushima Island, February 3, 1926.
All the samples were rich in pelagic diatoms, especially in
Coscinodiscus, Rhizosolenia, Chaetoceras, and Stephanopyxis.
Sample 1 contained Coscinodiscus plankton; sample 2, a Rhizo-
solenia-Chaetoceras plankton; sample 3 was very rich in Crus-
tacea and contained some Coscinodiscus, Eucampia, and Stepha-
nopyxis; sample 4 was the richest in pelagic forms, especially
of the genera Stephanopyxis, Lauderia, Baeteriastrum, and
Chaetoceras; sample 5, collected not far from Tsushima Island,
contained small Chaetoceras.
All these samples yielded a considerable number of diatoms,
which I have described and enumerated as a contribution to
the geographic distribution of this important group of marine
organisms. The present list contains the names of seventy
95
96 The Philippine Journal of Science 1931
forms, among which are some new to science. This note is
illustrated with drawings by the author. The number or num-
bers after the comment on each species correspond to the local-
ities given above.
COSCINODISCUS EXCENTRICUS Ehrenber*.
A. Schmidt, Atlas Diatom., pi. 58, figs. 46-49; V. Heurck, Synopsis
(1880-S5) pL 130, figs. 4, 7, and 8.
Diameter of valve 0.037 to 0.042 millimeter. Markings poly-
gonal, decreasing towards the border. Apiculi distinct. Geo-
graphic distribution: Atlantic and Pacific Oceans and the Sea
of Japan. Localities 1, 3, and 4.
COSCINODISCUS FRAGILISSIMUS Grunow.
Grunow in V. Heurck, Synopsis (1880-85) pi. 128, fig. 4.
Ethmodiscus convexus Castracane, Diat. Challenger Exped. (1886)
167, pi. 3, fig. 9.
Diameter 0.085 to 0.1 millimeter (in Castracane 0.129, in
Grunow 0.316 millimeter). Central space minute, indistinct.
Markings forming invisible striations and minute denticules,
scattered on the whole surface of the disk. Geographic dis-
tribution: Arafura Sea. Locality 1.
COSCINODISCUS CONCINNUS W. Smith. Plate 1, fiffs. 1-2, 4-6.
A. Schmidt, Atlas Diatom. (1886) pi. 114, figs. 8, 9; Rattray, Revi-
sion Coscin. (1889) 531; Peragallo, Diat. Mar. France (1897-1908)
pi. 115, fig. 12.
Disk covered with radiating lines of small granules. Diam-
eter of disk 0.13 to 0.37 millimeter. Radiating lines are sep-
arated by rows of very minute granules, which pass from as
many points, disappear towards the center or form a central
rosette. Several valves were found with abnormal areoles,
forming a rim in the middle part or two centers. Castracane's
Coscinodisctis papuanus, from New Guinea, and Coscinodiscus
mirificus belong to our species. Geographic distribution: At-
lantic and Pacific Oceans. Localities 1, 2, 3, and 4.
COSCINODISCUS RADIATUS Ehrenberg. Plate 1, fi*. 3.
A. Schmidt, Atlas Diatom, pi. 60, figs. 1-6, 9, 10; pi. 61, fig. 13; pi.
65, fig. 8; pi. 113, figs. 8, 21; Rattray, Revision Coscin. (1889) 514.
Markings four in 0.01 millimeter, gradually decreasing to-
wards the border. Geographic distribution: Atlantic and Pa-
cific Oceans, Sea of Japan, Hong Kong, Dairen. Localities 1, 2,
and 3.
46,1 Skvortzow: Diatoms of Korean Strait 97
PLANKTONIELLA SOL (Wallich) Schutt- Plate 9, fig:. 10.
Schutt, Pflanzenleb. d. Hochsee (1893) 20, fig. 8.
Coscinodiscus sol Waulich, Trans. Micr. Soc. 8 (1860) 38, figs. 1-2.
Cestodiscus sol Grunow in V. Heurck, Synopsis (1880-85) pi. 12, fig.
9; A. Schmidt, Atlas Diatom. (1878) pi. 58, figs. 41, 42, 45; Karsten,
Indisch. Phytopl. (1907) 369, pi. 39, figs. 1-12; Hustedt, Kieselal-
gen (1929) 465-67, fig. 259; Rattray, Revision Coscin. (1889) 466.
Valve flat, disklike. Length 0.052 to 0.12 millimeter. Mark-
ings distinct, in radiate rows, decreasing from the center
outward. The outer portion of the valve makes a broad,
scarcely siliceous disk. Geographic distribution: Common in
plankton of Atlantic and Pacific Oceans, Java Sea, Sea of Japan ;
fossil in Cambridge and Barbados deposits. Locality 4.
ACTINIPTYCHUS UNDULATUS (Bailey) Ralfs.
A. Schmidt, Atlas Diatom. (1874-81) pi. 1, figs. 1-4, 6, 8, 9; pi. 29,
figs. 4-8; pi. 109, fig. 1; pi. 122, figs. 1, 3; V. Heurck, Synopsis
(1880-85) pi. 122, figs. 1, 3.
Valve disklike. Length 0.025 to 0.09 millimeter. Areoles 3
in 0.01 millimeter. Geographic distribution: Atlantic and Pa-
cific Oceans, Sea of Japan, Dairen. Locality 4.
CORETHRON PELAGICUM Bran. Plate 8, &g. 14.
Schroder, Phytopl. warmer Meere (1906) 343, fig. 3; Hustedt, Kiesel-
algen (1929) 547, figs. 312a, b, c.
Cell robust, cylindrical, 0.095 to 0.115 millimeter in diameter,
with rounded ends and long spines on both sides. Spines very
delicate, smooth. Geographic distribution: Atlantic and Pa-
cific Oceans; Hong Kong. Locality 4.
EUCAMPIA ZODIACUS Ehrenbergr. Plate 2, figrs. 5, 6.
Kutzing, Bacillar. (1844) 143, pi. 21, fig. 21; W. Smith, Brit. Diatom.
(1853-56) 2, 25, pi. 35, fig. 299; Schutt, Bacillariales (1896) 89,
figs. 46A, 147B; Okamura, Littoral Diatoms Japan (1911) 6-7, pi.
11, figs. 33a~d; Peragallo, Diat. Mar. France (1897-1908) 376, pi.
95, fig. 2; V. Heurck, Synopsis (1880^85) pi. 95, figs. 17-18.
Cell elliptical forming curved chain. Foramina oval. Geo-
graphic distribution: Atlantic and Pacific Oceans; Malay Archi-
pelago; in Japanese waters known from Goza, Toshima, and
Boshyu. Localities 1, 2, 3, and 4.
EUCAMPIA BICONCAVA (Cleve) Ostenfeld. Plate 2, fig:. 9.
Ostenfeld, Flora Koh Chang, Mar. Plank. Diat. (1902) 23.
Eucampia hemiauloides Ostenfeld in Ostenfeld and Schmidt, Plankton
Rode Hav og Adenbugten (1901) 157-58, fig. 8.
259737 7
98 The Philippine Journal of Science 1931
Climacodium biconcavum Cleve, Phytoplank. (1897) 22, pi. 2, figs.
16, 17; Okamura, Littoral Diatoms Japan (1911) 8, pi. 11, fig. 35.
C. H. Ostenfeld gives the following description of this alga:
Chain straight, cells slightly siliceous, nearly as long as wide (length
0.04 to 0.06 millimeter, width 0.035 to 0.065) ; side view elliptic; front view
symmetrical on both sides of the longitudinal axis; processes of the valves
short; valves membraneous; connecting zone very finely annulated. Chro-
matophores numerous disciform.
Geographic distribution : Atlantic and Pacific Oceans, Mediter-
ranean and Red Seas, Gulf of Aden, Malay Archipelago; Sea
of Japan, Zenidzu. Locality 4.
STEPHANOPYXIS TURRIS (Greville and Arnott) Ralfs. Plate 2, fig. 4.
Pritchard, Infusor. (1861) 826, pi. 5, fig. 74; A. Schmidt, Atlas Dia-
tom. (1888) pi. 130, figs. 42, 43.
Stephanopyxis appendiculata Ehrenberg, in Microgeol. (1854) pi. 18,
fig. 4; Hustedt, Kieselalgen (1928) 304-306, fig. 140.
Cell cylindrical, with rounded ends, 0.03 to 0.045 millimeter
in breadth and 0.074 to 0.085 in length. The surface is densely
cellular. Geographic distribution: Atlantic and Pacific Oceans;
known from Japanese waters. Localities 3 and 4.
STEPHANOPYXIS PALMERIANA (Greville) Grunow. Plate 2, figs. 1, 2.
Grunow, Diatomeen Franz Josefs-Land (1884) 90.
Stephanopyxis var. javanica Grunow in A. Schmidt, Atlas Diatom.
pi. 130, fig. 44.
Stephanopyxis palmeriana var. japonica in Okamura, Littoral Diatoms
Japan (1911) 2, pi. 8, fig. 2; Gran and Yendo, Japan. Diatoms
(1914) 26-27.
Stephanopyxis campana Castracane, Diat. Challenger Exped. (1886)
88, pi. 19, fig. 14; Karsten, Indisch. Phytopl. (1907) pi. 54, figs. 9a,
b; Hustedt, Kieselalgen (1928) 308-9, figs. 147a^.
Cells cylindrical, 0.045 to 0.135 millimeter in diameter, forming
a long chain. Cells covered with areoles, large in the upper part
and small in the middle part of the cell. Geographic distribu-
tion: Atlantic and Pacific Oceans. Hong Kong; in Japanese
waters known from Shinojima, Shirahama, Goza, Mizaki, Misumi,
Yenoshima, Akashi Channel, and Yeddo Bay. Localities 3 and 4.
STEPHANOPYXIS PALMERIANA forma CURTA forma nov. Plate 2, fig. 3.
Valve flat, 0.105 to 0.12 millimeter broad and 0.04 to 0.045
in length. Localities 3 and 4.
THALASSIOSIRA HYALINA (Grunow) Gran. Plate 2, fig. 10.
Gran, Biblioth. Botan. 42 (1897) 4, pi. 1, figs. 17, 18.
Coscinodiscus hyalinus Grunow, Kongl. Sv. Vet. Akad. Handl. 17, No.
2 (1884) 113, pi. 7, fig. 128.
46,1 Skvortzow: Diatoms of Korean Strait 99
Thalassiosira clevei Gran, Norske Nordh. Exped. Bot. Protoph. (1897)
29, pi. 4, figs. 60-62; Peragallo, Diat. Mar. France (1908) 438, pi.
120, fig. 9.
Thalassiosira gravida Clevb in Okamura, Littoral Diatoms Japan
(1911) 2, pi. 8, fig. 3; Hustedt, Kieselalgen (1928) 323-24, fig.
159.
A pelagic diatom forming a long chain composed of flat cells.
Our plant has cells 0.045 to 0.052 millimeter in diameter. Ac-
cording to J. Rattray (1889) :
Diameter of the valve is 0.025 millimeter. Central space minute, incon-
spicuous, bearing isolated puncta. Markings punctiform, subequal, 24 in
0.01 millimeter, rows radial to subparallel in inconspicuous fasciculi; api-
culi numerous, distinct, in a single circlet. Border broad, hyaline.
Geographic distribution: Atlantic and Pacific Oceans. Local-
ity 4.
LAUDERIA BOREALIS Gran. Plate 2, fig. 11.
Gran, Nyt. Mag. f. Naturvid. 38 (1900) 110, pi. 2, figs. 5-9.
Lauderia annulata Cleve, Diatoms of Sea of Java (1873) 8, pi. 1,
fig. 7.
Lauderia compressa Peragallo, Diatom. Mar. France (1897-1908) pi.
121, fig. 2.
Thalassiosira nordenskioldii Cleve in Okamura, Littoral Diatoms Ja-
pan (1911) 2, pi. 8, fig. 4; Hustedt, Kieselalgen (1928) 549-50.
Chain composed of cylindrical f rustule, orbicular in side view,
near the margin covered with numerous short, hairlike spines.
Sculpture consists of very fine puncta. Geographic distribu-
tion: Atlantic and Pacific Oceans; in Japanese waters known
from Shirahama. Localities 1, 3, and 4.
SCHROEDERELLA DELICATULA (Peragallo) Pavillard. Plate 2, fig. 13.
Pavillard, Obser. Diat. (1913) 60, 126.
Lauderia delicatida Peragallo, Bull. Soc. Hist. Nat. Toulouse (1888)
22, 81, pi. 6, fig. 46.
Lauderia delicatula Peragallo, Diat. Mar. France (1897-1908) pi.
121, figs. 4, 8, 9.
Lauderiopsis costata Ostenfeld, in Ostenfeld and Schmidt, Plankton
Rode Hav. og Adenbugten (1901) 158-59, fig. 10.
Detonida schroederi Gran, Nord. Plankton (1906) 22; Hustedt in A.
Schmidt, Atlas Diatom. (1920) pi. 320, figs. 16-17;, pi. 321, fig. 4;
Kieselalgen (1929) 551-53, fig. 314.
Chain straight, composed of many cylindrical cells 0.025 to
0.04 millimeter broad, rectangular in front view. Geographic
distribution: Common in Atlantic and Pacific Oceans. Local-
ity 4.
100 The Philippine Journal of Science 1931
LEPTOCYLINDRUS CURVATUS sp. nov. Plate 2, fi*. 14.
Cell cylindrical, rectangular in front view, 0.004 to 0.0045 mil-
limeter broad, 0.007 to 0.008 in length. Chromatophores nu-
merous. Locality 4.
DITYLIUM BRIGHTWELLII (West) Grunow. Plate 2, figs. 7, 8.
V. Heurck, Synopsis (1880-85) pi. 114, figs. 3-9; Traite Diatom.
(1889) 424, pi. 17, %. 606; A. Schmidt, Atlas Diatom, pi. 152, figs.
10-13; Peragallo, Diatom. Mar. France (1897-1908) 395-96, pi.
96, figs. 6-11; Schroder, Phytopl. Antarkt. Meeres (1906) 353-55,
figs. 22a-e.
Ditylium sol V. Heurck in Okamura, Littoral Diatoms Japan (1911)
8, pi. 11, fig. 37.
A plankton species with a peculiar triangular valve with two
long horns from both sides. Geographic distribution : Atlantic
and Pacific Oceans ; in the Sea of Japan known from Tateyama,
Shirahama, Shima, Misaki, and Hong Kong. Localities 2 and 4.
CHAETOCERAS LORENZIANUM Grunow. Plate 3, fig. 4.
Grunow, Osterreich. Diatom. (1864) 157, pi. 5, fig. 13;, V. Heurck,
Synopsis (1880-85) pi. 82, fig. 2; Cleve, Diatom. Arctic Sea (1897)
21, pi. 1, figs. 13-15; Gran, Nord. Plankton (1906) 76, fig. 90; Oka-
mura, Chaetoceras and Peragallia (1907) 93, pi. 4, figs. 38-39; Lit-
toral Diatoms Japan (1911) 7, pi. 11, fig. 31.
Chaetoceras celluloswm Lauder, Diatom. Hong Kong (1864) 78, pi. 8,
fig. 12; Gran and Yendo, Japan. Diatoms (1914) 9; Peragallo,
Diatom. Mar. France (1897-1908) 484, pi. 131, figs. 1-3; Hustedt
in A. Schmidt, Atlas Diatom. (1920) pi. 321, figs. 18-19; pi. 322,
%. 1.
Cells forming a straight chain 0.008 to 0.012 millimeter broad.
Cell rectangular with projecting angles. Foramina large, broad
elliptic or quadrangular with round angles. Setae thin, their
basal parts almost parallel to the chain axis. Terminal setae
rather well differentiated and disposed parallel or more or
less divergent. Geographic distribution: Atlantic and Pacific
Oceans; in Japanese waters known from Boshu, Misaki, Eno-
shima, Akashi, and Formosa Channel. Localities 2, 4, and 5.
CHAETOCERAS JAVANICUM Clere. Plate 3, fig. 2.
Cleve, Diatoms Sea of Java (1873) 11, pi. 2, fig. 13; Peragallo, Dia-
tom. Mar. France (1897-1908) 480, pi. 130, figs. 1, 2; Hustedt in
A. Schmidt, Atlas Diatom. (1920) pi. 323, figs. 1, 2.
Cells forming a straight chain 0.02 to 0.04 millimeter broad.
From the front view the cell is rectangular with projecting
angles. Foramina large, broad elliptic; setae thin, with their
basal parts almost parallel to the chain axis. Terminal setae
parallel or more or less divergent. Chromatophore single.
46,1 Skvortzow: Diatoms of Korean Strait 1Q1
Geographic distribution: Indian Ocean and Sea of Japan. Lo-
calities 2, 4, and 5.
CHAETOCERAS SIAMENSE Ostenfeld. Plate 3, fig. 3.
Ostenfeld, Flora Koh Chang (1902) 21, fig. 17.
Chaetoceras lauderi Ralfs var. in Lauder, Diatom. Hong Kong* (1864)
77, pi. 8, fig. 3.
Chaetoceras misumense Gran and Yendo, Japan. Diatoms (1914)
14-15, fig. 7;, Ikari, Chaetoceras Japan. (1928) 257-58, fig. 11.
I quote the diagnosis of Gran and Yendo, as follows :
The frustules are quadrangular, measuring 0.02 to 0.03 millimeter in
breadth, and 0.02 to 0.04 millimeter in height, and are elliptical in a valvar
view. The setae spring directly from the corners of the valves, at first di-
verging and then gradually bending in a direction parallel to the axis of
the chain. Their terminal half is armed with minute spinous processes
arranged spirally. The terminal horns have a similar direction to that
of the setae, but bent more abruptly near the points of insertion and
then run almost straight out, forming an acute angle with one another.
The two horns are in one plane. They are more robust than the setae, and
like them are beset with minute processes. The foramen is elliptical or
broadly lanceolate in a surface view, but practically narrow lanceolate
in an optical section. There are two peculiar depressions in the middle
of each valve, as shown by Lauder. The girdle-bands become much nar-
rowed about the middle of the complanate side of the frustules. The rest-
ing spores have pallisade spines on the margins of both the primary
and the secondary valve. The primary valve is almost hemispherical, and
has numerous short spines over its whole surface; the secondary valve is
nearly similar, but is somewhat humped and has short spines condensed
about the summit.
The Korean specimens were without spores. Geographic dis-
tribution: Sea of Japan (Kushimoto, Seto, Misumi), South China
Sea, Hong Kong. Localities 2 and 5.
CHAETOCERAS BOREALE Bailey. Plate 3, fig. 1.
Bailey, Notes Microscop. Organ. (1854) 8, figs. 22-23.
Chaetoceras boreale var. brightwellii Cleve, Diat. Arctic Sea (1873)
12, fig. la.
Chaetoceras boreale Cleve, Treatise Phytopl. N. Atlantic (1897) 20,
pi. 1, %. 1.
Chaetoceras boreale var. brightwellii Cleve, Treatise Phytopl. N. At-
lantic (1897) 20, pi. 1, fig. 2; Peragallo, Diatom. Mar. France
(1897-1908) 476-77, pi. 127, figs. 2, 3; Okamura, Chaetoceras and
Peragallia Japan (1907) 90, pi. 3, figs. 18-20; Gran, Diatom. Arkt.
Meere (1904) 533, fig. 5; Nord. Plankton (1906) 73, fig. 87; Gran
and Yendo, Japan Diatom. (1914) 7; Hustedt in A. Schmidt, Atlas
Diatom. (1920) pi. 325, figs. 5, 6.
Chain straight, 0.025 to 0.04 millimeter broad. Cells in front
view quadrangular with rounded angles, shorter than the
102 The Philippine Journal of Science 1931
breadth. Girdle band attains over one-third of the cell height.
Valve convex. Foramina small and rhomboidal. Setae start di-
rect from these cones and each then coalesces with that of the
neighboring cell within the lateral sides of the chain. Setae
adorned with spines. Geographic distribution: Common in
northern seas, Atlantic and Pacific Oceans; known from Jap-
anese waters from Tateyama and Misaki. Localities 2, 3, and 4.
CHAETOCERAS REICHELTI Hustedt. Plate 7, &g. 2.
Hustedt in A. Schmidt, Atlas Diatom. (1921) pi. 344, fig. 6.
Chain straight, composed of 3 or 4 cells, 0.02 to 0.03 milli-
meter broad. Cells in front view quadrangular, twice longer
than breadth. Girdle band about one-half of the mantle height.
Setae long, punctate, issuing from the angles of the valves, cross-
ing each other, leaving short basal parts and diverging at an
obtuse angle. Terminal setae nearly parallel to the chain axis
or somewhat divergent from each other. Geographic distribu-
tion: See Adler-Hafen collected by Cohn in 1912. Locality 4.
CHAETOCERAS IKARI sp. nov. Plate 7, fig. 1.
Chain straight 0.008 to 0.015 millimeter broad, composed of
five to twenty cells. Frustules rectangular elongate, three to
four times as long as the breadth, with sharp projecting angles.
Valve concave or sometimes flat. Foramina large, broad, ellip-
tic. Setae thin, issuing from the angles of the valves, diverging
at an obtuse angle. Chromatophore single. Named in honor
of the well-known Japanese diatomist I. Ikari, Seto Biological
Station, Japan. Localities 2 and 4.
CHAETOCERAS SOCIALE Lauder. Plate 5, fig. 7.
Lauder, Diatom. Hong Kong* (1864) 77, pi. 8, fig. 1; Cleve, Diatoms
Baffin Bay (1896) 9, pi. 2, fig. 9; Gran, Baeillar. Karajakfjord
(1897) 26, pi. 4, fig. 54; Nord. Plankton (1906) 96, fig. 123; Gran
and Yendo, Japan. Diatoms (1914) 24; Okamura, Littoral Diatoms
Japan (1911) 7, pi. 11, fig. 30; Peragallo, Diatom. Mar. France
(1897-1908) 490, pi. 132, figs. 1-3.
Cells slender, aggregated, embedded in gelatine. Chain com-
posed of three to five cells. Valves broadly oval and flat, in
front view rectangular. Setae very delicate, thin, straight,
curved or undulated. The chromatophore single. Geographic
distribution: Arctic Sea, Atlantic and Pacific Oceans; known
from Japanese waters from Boshu Province, Volcano Bay, Mi-
4<3, i Skvortzow: Diatoms of Korean Strait IQ3
saki, Otaru Bay, Euoshima, and Akashi. Localities 1, 2, 4, and
5.
CHAETOCERAS PROTUBERANS Lauder. Plate 5, fi*. 4.
Lauder, Diatom. Hong Kong (1864) 79, pi. 8, fig. 11.
Chaetoceras didymum in Okamura, Chaetoceras and Peragallia (1907)
95, pi. 4, fig. 45a; Gran and Yendo, Japan. Diatoms (1914) 12-13,
fig. 5; Hustedt in A. Schmidt, Atlas Diatom. (1920) pi. 326, figs!
1, 5.
Cells forming a long chain, 0.005 to 0.008 millimeter broad.
Cell oblong with mamillif orm protuberance. Setae long, spinose,
coalesced. Terminal setse well developed, curved, with spirally
disposed punctations. Geographic distribution: Atlantic and
Pacific Oceans ; in Japanese waters known from Shima Province,
Misaki, Otaru Bay, and Misumi. Localities 2, 4, and 5.
CHAETOCERAS DIDYMUM Ehrenberg var. ANGLICA Gran. Plate 5, fig. 6.
Gran, Nord. Plankton, (1906) 80, fig. 95; Okamura, Chaetoceras and
Peragallia (1907) 95, pi. 4, figs. 44-47, excl. fig. 45«; V. Heurck,
Synopsis (1880-85) pi. 82, fig. 3.
Chaetoceras didymum, var. longicruris Cleve, Phytopl. (1897) 21, pi.
1, fig. 11; Peragallo, Diatom. Mar. France (1897-1908) 481, pi. 128,
fig. 3; Hustedt in A. Schmidt, Atlas Diatom. (1920) pi. 326, figs.
3, 4.
Cells forming a straight chain, 0.025 to 0.03 millimeter broad,
sometimes composed of twenty or more cells. Cells quadran-
gular or elliptical, convex in the middle, forming a large dot.
Setae arise from within the angles curved. Geographic distribu-
tion: In Japanese waters found in Shima Province, Misaki, Mi-
sumi, Enoshima, and Formosa Channel. Localities 2, 4, and 5.
CHAETOCERAS DIDYMUM Ekrenberg var. GENUINA Gran. Plate 5, figs. 3 and 5.
Gran, Nord. Plankton (1906) 79, fig. 94; Gran and Yendo, Japan.
Diatoms (1914) 12-13; Okamura, Chaetoceras and Peragallia (1907)
95, pi. 4, figs. 44-47, excl. fig. 45a; Hustedt in A. Schmidt, Atlas
Diatom. (1920) pi. 326, figs. 2, 7; Peragallo, Diatom. Mar. France
(1897-1908) 480-81, pi. 128, figs. 1, 2.
Cells forming a straight chain 0.03 to 0.055 millimeter broad,
composed of many cells. Cells broad elliptical, convex in the
middle. Setse starting from the angles of the valve, crossing
one another close to their insertions, diverging at an obtuse
angle. Terminal setse well developed, robute, straight. Geogra-
phic distribution: Atlantic and Pacific Oceans; known from
104 The Philippine Journal of Science 1931
Japanese waters from Tosa Province, Misumi, Enoshima, and
Akashi. Localities 2, 3, 4, and 5.
CHAETOCERAS RADIANS Schtttt. Plate 5, fig. 2.
Schutt, Chaetoceras und Peragallia (1895) 41, fig's. 10a-c; Peragallo,
Diatom. Mar. France (1897-1908) 490-91, pi. 133, fig. 4; Gran,
North Polar Exped. (1900) 26; Nord. Plankton (1906) 97, fig. 124.
Cell forming a spiral and curved chain, 0.008 to 0.01 mil-
limeter broad. Cell small in front view, rectangular. Fora-
mina oblong or hexagonal. Setae only on one side, straight, thin.
Chromatophore single. Geographic distribution: Atlantic and
Pacific Oceans. Localities 4 and 5.
CHAETOCERAS DECIPIENS Cleve. Plate 6, figs. 3 and 4.
Clbve, Diatoms Arctic Sea (1873) 11, fig. 5; Gran, Protoph., Diatom.,
etc. (1897) 13, pi. 1, figs. 2, 3; pi. 3, figs. 3, 4; Diatom. Arkt. Meere
(1904) 535-38, pi. 17, figs. 1-6; Gran and Yendo, Japan. Diatoms
(1914) 8-9, figs. 3a, b; Peragallo, Diatom. Mar. France (1897-1908)
485, pi. 131, figs. 4-8.
Chaetoceras grunowii Schutt, Chaetoceras und Peragallia (1895) 43,
figs. 14a, 6.
Cells forming a straight chain, rectangular in a broader front,
with round or somewhat projecting angles. Valves broad ellip-
tic, convex in the middle. Foramina elongate, slightly con-
stricted in the middle. Basal parts of the setae parallel to chain
axis, then parallel or diverging at an obtuse angle. Terminal
setae diverging in a variable angle. The terminal and the lat-
eral setae are clearly punctated, but in some chains such mark-
ings are entirely lacking. Chromatophores numerous. Geo-
graphic distribution: Very common in Atlantic and Pacific
Oceans; in Japanese waters found at Otaru Bay, Echigo Prov-
ince, Misumi, and Enoshima. Localities 2, 4, and 5.
CHAETOCERAS COMPRESSUM Lander. Plate 5, fig. 1.
Lauder, Diatom. Hong Kong (1864) 78, pi. 8, fig. 6; Cleve, Plankton.
Ciliof. och. Diatom. (1894) 12, pi. 2, fig. 3; Schutt, Chaetoceras und
Peragallia (1895) 43, figs. 16, a, b; Ostenfeld, Flora Koh Chang
(1902) 94, pi. 3, figs. 8-11.
Chaetoceras contortum Schutt, Diatom. Chaetoceras (1888) pi. 3,
fig. 4.
Chaetoceras medium Schutt, Chaetoceras und Peragallia (1895) 43,
fig. 15;, Gran, Bacillar. Karajakfjord. (1897) 14, pi. 2, fig. 32; Gran
and Yendo, Japan. Diatoms (1914) 10, figs. 4a-d; Peragallo,
Diatom. Mar. France (1897-1908) 488-89, pi. 134, fig. 8.
Chain straight or slightly curved when composed of many cells,
0.016 to 0.024 millimeter broad. Foramina oblong or constricted
46,1 Skvortzow: Diatoms of Korean Strait 105
in the middle part. Lateral view of valve compressed oval.
Sete arise from a little within the angles, long, robust, with
strong undulations and with verrucose dots small and very thin.
Geographic distribution : A well-distributed species known from
the Sea of Japan, Volcano Bay, Echigo Province, Misaki, Yeddo
Bay, Misumi, Tateyama, Enoshima, Akashi, and Formosa Chan-
nel. Localities 2, 4, and 5.
CHAETOCERAS TORTISSIMUM Gran. Plate 6, fi«r. 2.
Gran, Nord. Plankton (1906) 95-96, fig. 122; Pavillard, Danish Ocean.
Exped. (1925) 52, fig. 87; Ikari, Chaetoceras of Japan (1928)
532-33, figs. 5a, b.
Chain straight or curved. Cells in front view rectangular,
with some rounded angles. Setae delicate, thin, smooth, most
perpendicular to the chain axis. Chromatophores solitary.
Geographic distribution: Atlantic and Pacific Oceans; Sea of
Japan at Seto and Oshoro. Locality 5.
CHAETOCERAS ATLANTICUM Cleve. Plate 7, figs. 3-5.
Cleve, Diat. Arctic Sea (1873) 11, pi. 2, fig. 8; Gran, Nord. Plankton
(1906) 64, fig. 74; Gran and Yendo, Japan. Diatoms (1914) 3-5, fig.
1; Kabsten, Phytopl. Antark. Meere (1905-6) 115, pi. 15, fig. 9;
Okamura, Chaetoceras and Peragallia (1907) 89, pi. 4, figs. 56-62.
Chaetoceras atlanticum var. tumescens Grunow in V. Heurck. Sy-
nopsis (1880-85) pi. 81, fig. 6.
Chaetoceras dispar Castracane, Diatom. Challenger Exped. (1886)
76, pi. 8, fig. 6.
Chaetoceras compactum Schutt, Chaetoceras und Peragallia (1895)
46, fig. 23.
Chaetoceras skeleton Schroder, Phytopl. warm. Meere (1906) 337.
Chain straight and composed of many cells. Cells in front
view quadrangular or, frequently, much compressed; shorter
than the breadth. Girdle band about one-third of the mantle
height. Setae long, smooth, curved. Geographic distribution:
Atlantic and Pacific Oceans; in Japanese waters known from
Tosa, Kuriles, Enoshima, Volcano Bay, and Otaru Bay. Local-
ity 4.
CHAETOCERAS DADAYI Pavillard. Plate 6, fig. 1.
Pavillard, Observ. Diatoms (1913) 131-33, fig. 2; Danish Ocean. Ex-
ped. (1925) 41, figs. 546; Ikari, Chaetoceras of Japan (1926) 519-
20, figs. 2c, d.
Chain straight, composed of five or more cells, 0.025 to 0.04
millimeter broad. Frustules in front view rectangular with
rounded angles. Foramina very narrow or indistinct. Setae
106 The Philippine Journal of Science 1931
issuing from the margin, long and thickened, with densely dis-
posed spines. Terminal setse not differentiated. Chromato-
phores small, numerous, and passing into the setae. Geographic
distribution: Atlantic and Pacific Oceans, Mediterranean Sea,
Sea of Japan, Seto. Locality 4.
CHAETOCERAS PERUVIANUM BrightwelL Plate 4, figs. 4, 5.
Brightwell, Filam. longhorned Diatom. (1856) 107, figs. 16, 17; Cleve,
Diatoms of Sea of Java (1873) 8, pi. 2, fig. 8; Karsten, Phytopl.
Antark. Meere (1905) 166, pi. 31, fig. 4; Okamura, Chaetoceras and
Peragallia (1907) 91, pi. 4, figs. 67-75; Peragallo, Diatom. Mar.
France (1897-1908) 475, pi. 125, fig. 1.
Peragallia meridiana Schutt, Chaetoceras und Peragallia (1895) 48,
figs. 28a, b.
Cells solitary, or forming a very short filament composed of
two to four cells, 0.015 to 0.025 millimeter broad. Cells from
front view quadrangular or elongated with rounded angles. The
elongated cells have a transversal costation, sometimes zigzag
in the middle part. Valve convex. Setse very robust, straight,
curved, covered with solid spines and transversely very thinly
striated. Chromatophores small, round, numerous. A chain-
forming Peragallia meridiana Schutt, Iiro Ikari describes as
Chaetoceras okamurai Ikari. Geographic distribution: Atlan-
tic, Indian, and Pacific Oceans; in Japanese waters found in
Tosa Province, Shima Province, Misaki, Enoshima, Akashi, and
Seto. Localities 2, 4, and 5.
CHAETOCERAS SALTANS Cleve. Plate 4, fi*. 3.
Cleve, Phytopl. N. Atlantic (1897) 22, pi. 1, fig. 8; Peragallo, Diatom,
Mar. France (1897-1908) 476, pi. 126, fig. 1.
Cells solitary, 0.015 to 0.018 millimeter broad, from the front
view quadrangular, with rounded angles. Girdle band narrow.
Setae robust and spinulose. Related to Chaetoceras perwvianum
Brightw. and somewhat to Chaetoceras criophilum f. volans
(Schutt) Gran. Geographic distribution: Atlantic and Pacific
Oceans. Locality 4.
CHAETOCERAS AFFINE Lauder. Plate 4, fig. 2.
Lauder, Diatom. Hong Kong (1864) 78, pi. 8, fig. 5.
Chaetoceras ralfsii Cleve, Diatoms of Sea of Java (1873) 10, pi. 3,
fig. 15.
Chaetoceras ralfsii Cleve in Karsten, Phytopl. Antarkt. Meere (1906)
168-69, pi. 33, figs. 17-18.
Chaetoceras schuttii Cleve, Plankton., Ciliof. och Diatom. (1894) 14,
pl. 1, fig. 1.
Chaetoceras distichum Schutt, Chaetoceras und Peragallia (1895)
37, figs. 2a, 6.
46,1 Skvortzow: Diatoms of Korean Strait 107
Chaetoceras angulatum Schutt, Chaetoceras und Peragallia (1895)
37, figs. la-d.
Chaetoceras procerum Schutt, Chaetoceras und Peragallia (1895)
38, fig. 3a, b; Peragallo, Diatom. Mar. France (1895-1908) 478-
79, pi. 129, fig. 3.
Cells forming a straight chain, 0.015 to 0.024 millimeter broad,
composed of five to fifteen cells. Frustules in a broad front
quadrangular with somewhat pointed angles. Foramina nar-
rowly lanceolate, slightly constricted in the middle. Setae all
alike, smooth, disposed in a valval view at about right angles
with one another. Terminal setae well developed, curved, horn-
like, with minute elevations spirally disposed. Chromatophore
is one large plate, parietally in each cell. Resting spores 0.018
to 0.03 millimeter in diameter, primary valve nearly hemispher-
ical with short spines all over the surface. Secondary valve
greatly convex with long spines on the middle part.
According to Gran and Yendo, Chaetoceras affine is extremely
variable in its form and in Japanese waters two types were
found, one with rectangular, another with narrow, subcylin-
drical frustules. The terminal horns are less curved or diver-
gent. Geographic distribution: Chaetoceras affine is widely
distributed in the warmer parts of the Atlantic, Indian, and
Pacific Oceans; in Japanese waters it has been recorded at
Boshu Province, Shima Province, Enoshima, Akashi, Yeddo Bay,
and Misaki. Localities 2, 4, and 5.
CHAETOCERAS MESSANENSE Castracane. Plate 4, fig. 1.
Castracane, Contrib. Fl. Mediter. (1875) 32, pi. 1, fig. 1.
Chaetoceras sp. in Lauder, Diatom. Hong Kong (1864) pi. 3, fig. 8.
Chaetoceras furca Cleve, Phytopl. N. Atlantic (1897) 21, pi. 1, fig. 10.
Chaetoceras furca Cleve in Karsten, Phytopl. Antark. Meere (1905)
169, pi. 32, figs. 13 a, b.
Chaetoceras furca Cleve var. macroceras Schroder in Okamura, Chae-
toceras and Peragallia (1907) 99, pi. 3, fig. 7; Gran, Nord. Plank-
ton (1906) 87, fig. 108; Peragallo, Diatom. Mar. France (1897-
1908) 488, pi. 129, fig. 1; Hustedt in A. Schmidt, Atlas Diatom.
(1920) pi. 322, figs. 4 and 7; pi. 325, fig. 3.
Chain straight, 0.011 to 0.028 millimeter wide and 0.1 to 0.15
in length. Cells in front view rectangular with projecting an-
gles, valves concave, foramina lanceolate. Girdle band rather
longer than about a half of the cell height. Setae of two types.
One type is thin, short, starting from the angles of the valve,
crossing one another close to their insertions, diverging at an
obtuse angle. The other is robust, furcate at the end with
108 The Philippine Journal of Science 1931
spirally disposed punctations. Terminal setae not differentiated,
short, simple. Chromatophore solitary. Locality 5.
BACTERIASTRUM VARIANS Lauder. Plate 8, figs. 1, 3, 5-7.
Lauder, Diatom. Hong Kong (1864) 6, figs. 1-5; Karsten, Phytopl.
Antark. Meere (1905) 170, pi. 34, fig. 1; Peragallo, Diatom. Mar.
France (1897-1908) 470, pi. 136, fig. 1-5.
Chaetoceras varians (Lauder) V. Heurck, Synopsis (1880-85) 195,
pi. 70, figs. 3-5.
Bacteriastrum spirillum Castracane, Diatom. Challenger Exped.
(1886) 83, pi. 19, fig. 2; Hustedt in A. Schmidt, Atlas Diatom.
(1920) pi. 328, figs. 1-5 and 11; Ikari, Bacteriastrum of Japan
(1927) 421-22, fig. 1.
Chain straight, composed of many cells. Frustules cylin-
drical, with ten to fourteen furcated setae covered with thin
undulations. Terminal horns ten, curved at the ends and cov-
ered with spiral markings. Geographic distribution: Atlantic
and Pacific Oceans ; known from the Japanese waters of Oshoro,
Hakodate, Seto, Kushimoto, Takashima, and Goza. Localities
2, 3, and 5.
BACTERIASTRUM HYALINUM Lauder. Plate 8, fig. 2.
Lauder, Diatom. Hong Kong (1864) 6, pi. 3, figs, la, 6.
Bacteriastrum spirillum Castracane, Diatom. Challenger Exped. (1886)
83, pi. 24, fig. 1.
Bacteriastrum varians var. hyalina Lauder in Peragallo, Diatom. Mar.
France (1897-1908) 470, pi. 136, fig. 6; Pavillard, Observ. Diatoms
(1916) 27, pi. 1, fig. 4; Danish Oceanogr. Exped. (1925) 37, fig.
58; Ikari, Bacteriastrum of Japan (1927) 422-23, fig. 2.
According to Ikari this species has —
Chain straight and composed of many cells. Foramina not very large.
Length of the cell is about equal to, but generally a little shorter than
the breadth (diameter 0.013 to 0.056 millimeter). Setae, from 7 to 25
in number. Intermediate ones unite one another to form radial rays of
nearly equal length to or at least a half of the diameter of the cell. The
mode of bifurcation is greatly different from the preceding species; the
plane which involves the furcated parts, is placed parallel to the chain
axis, giving a very spiny appearance to the chain. The terminal horns
are gradually curved and bent downwards, and show spiral undulations on
them. Chromatophores numerous and small, the resting nucleus is sit-
uated in the center of the cell.
Geographic distribution: Atlantic and Pacific Oceans, Sea of
Japan, Seto, Kushimoto, Enoshima, and Akashi. Localities 2,
4, and 5.
4<J'1 Skvortzow: Diatoms of Korean Strait 1Q9
BACTERIASTRUM MINUS Karsten. Plate 8, fi*. 4.
Kabsten, Phytopl. Antark. Meere (1905) 171, pi. 33, fig. 21.
Bacteriastrum hyalinum Lauder in Okamura, Littoral Diatoms Japan
(1911) 6, pi. 10, fig. 28; Ikari, Bacteriastrum of Japan (1927)
426-27, fig. 6.
Chain straight, composed of many cells. Frustules disklike,
0.018 to 0.025 millimeter in diameter. Intermediate and ter-
minal setae short and delicate, directed obliquely outwards to the
chain axis. Geographic distribution: Atlantic and Pacific
Oceans; in Japanese waters known from Oshoro, Hakodate, Ta-
teyama, Kashimoto, Seto, Takashima, and Goza. Locality 4.
BACTERIASTRUM COMOSUM Pavillard var. HISPIDA (Castracane) Ikari. Plate 8, %. 8.
Ikari, Bacteriastrum of Japan (1927) 428-29, fig. 86.
Bacteriastrum wallichi Ralfs. var. hispida Castracane, Diatom. Chal-
lenger Exped. (1886) 83, pi. 29, fig. 6.
Bacteriastrum varians Lauder var. hispida (Castr.) Schroder in
Schroder, Phytopl. Warm. Meere (1906) 347, fig. 11.
Bacteriastrum varians var. hispida in Okamura, Littoral Diatoms Ja-
pan (1911) 7, pi. 10, figs. 29 f-g; Hustedt in A. Schmidt, Atlas
Diatom. (1920) pi. 328, fig. 12.
Chain straight, composed of many cells. Frustules cylindrical,
valve flat. Intermediate setae long, furcate, curved. Terminal
setae distinct, robust, hornlike, undulated and covered with short
spines. Geographic distribution: Atlantic and Pacific Oceans,
Sea of Japan, Seto, and Kushimoto. Localities 2 and 4.
BIDDULPHIA SINENSIS Greville. Plate 8, fig. 9.
Greville, Trans. Micr. Soc. (1866) 81, pi. 9, fig. 16; Ostenfeld and
Schmidt, Plank. Rode Hav og Adenbugten (1901) 152, fig. 6;* Flora
Koh Chang (1902) 25, fig. 21; Cleve, Diatoms of Sea of Java (1873)
6; A. Schmidt, Atlas Diatom. (1888) pi. 122, figs. 22, 23, 24; Leud-
uger-Fgrtmorel, Diatomees Malaisie (1892) 39.
A large pelagic Biddulphia with very fine striation. Valve
robust, 0.2 to 0.27 millimeter broad and 0.5 to 0.7 millimeter in
length. Geographic distribution: Red Sea, Malay Archipelago,
Java Sea, South China Sea, Norway, and Sea of Japan. Lo-
cality 4.
BIDDULPHIA AURITA Brebisson var. ORIENTALIS Mereschkowsky.
Mereschkowsky, Polynesian Diatoms (190O-1902) 119; A, Schmidt,
Atlas Diatom, pi. 120, figs. 5, 6; Skvortzow, Marine Diatoms Dairen
(1929) 420; Marine Diatoms Siberian Shore (1929) 59, fig. 15.
Frustules without spines, 0.042 to 0.051 millimeter in length
and 0.018 to 0.037 in breadth. Geographic distribution: Com-
110 The Philippine Journal of Science 1931
mon in littoral zone in Pacific and Atlantic Oceans, Polynesian,
Sea of Japan, and Bay of Bengal. Localities 1 and 2.
BIDDULPHIA PULCHELLA Gray. Plate 2, fig:. 12.
W. Smith, Brit. Diatom. (1853-56) pi. 44, fig. 321; A. Schmidt, Atlas
Diatom., pi. 118, figs. 26-33; V. Heurck, Synopsis (1880-85) pi.
97, figs. 1-3; Peragallo, Diatom. Mar. France (1897-1908) 376-77,
pi. 93, figs. 1, 2.
Biddulphia biddulphiana (Smith) Boyer in Okamura, Littoral Dia-
toms Japan (1911) 9, pi. 12, fig. 42.
A cosmopolitan diatom, forming a straight chain composed of
robust triangular frustules. The length and the breadth of the
cells vary from 0.045 to 0.105 millimeter. Geographic distribu-
tion: Common in the littoral zone and sometimes in plankton.
Atlantic and Pacific Oceans ; environs of Vladivostok and Dairen.
Localities 1 and 2.
BIDDULPHIA LONGICORNIS Greville. Plate 2, fig. 16.
A. Schmidt, Atlas Diatom, pi. 118, fig. 10; Okamura, Littoral Diatoms
Japan (1911) 11, pi. 12, fig. 46.
Valve seen in face view broadly elliptical, with pointed ends
forming an obtuse angle, in which, close to the apices, arise two
long horns. Valve ornamented with rows of beads running lon-
gitudinally along the valve. In side or girdle view each valve
shows two very long outwardly curving horns and two long
spines in the middle part of the ends. Girdle broad and curved
with closely set rows of transverse beading. Length of the valve
0.066 to 0.075 millimeter, breadth 0.074 to 0.085 millimeter.
Geographic distribution: Pacific Ocean. Locality 4.
THALASSIOTHRIX ANTARCTICA Schimper forma JAPONICA forma nov. Plate 9, figs.
1 and 2.
Valve straight, linear, apices slightly produced. Length, 1.6
to 1.95 millimeters ; breadth, 0.0055 to 0.0075. Striae fifteen in
0.01 millimeter. Our form differs from the typical T. antarc-
tica by its straight valve. Geographic distribution : Atlantic and
Pacific Oceans. Locality 4.
THALASSIOTHRIX NITZSCHIOIDES Grunow. Plate 8, figs. 10 and 11.
V. Heurck, Synopsis (1880-85) pi. 43, fig. 7.
Synedra nitzschioides Grunow, 5sterreich. Diatom. (1862) 403, pi. 5,
fig. 18.
Thalassiothrix curvata Castracane, Diatom. Challenger Exped. (1886)
55, pi. 24, fig. 6.
Thalassiothrix frauenfeldi Cleve, Plankton., Ciliof., och Diatom. (1894)
6.
46,1 Skvortzow: Diatoms of Korean Strait HI
Thalassiothrix frauenfeldi var. nitzschioides in Jorgensen, Protophyten
und Protozoen Plankt. (1900) 20; Peragallo, Diatom. Mar. France
(1897-1908) 320, pi. 131, figs. 17, 18.
Cells forming star-shaped or zigzag clusters. Cells linear,
0.037 to 0.06 millimeter in length, 0.0025 to 0.006 in breadth;
striae 11 in 0.01 millimeter. Geographic distribution: Atlantic
and Pacific Oceans; known from Japan, Eastern and South
China Seas. Localities 2, 4, and 5.
THALASSIOTHRIX NITZSCHIOIDES var. JAVANICA Grunow. Plate 9, fig. 6.
V. Heurck, Synopsis (1880-85) pi. 98, figs. 11, 12.
Cells lanceolate, with slightly elongated, rounded apices.
Length, 0.042 to 0.055 millimeter; breadth, 0.0042 to 0.055;
striae marginal, 9 to 10 in 0.01 millimeter. Geographic distribu-
tion: Java Sea. Locality 4.
THALASSIOTHRIX FRAUENFELDII Grunow. Plate 9, fig. 8.
Cleve and Grunow, Arkt. Diatom. (1880) 109.
Asterionella frauenfeldii Grunow, Verhandl. Zool.-Bot. Gesellsch.
(1863) 140, pi. 14.
Asterionella frauenfeldii Grunow, Novara Algen (1867) 4.
Asterionella synedraeformis Greville, Ann. Nat. Hist. (1865) 4, pi.
5, figs. 5, 6.
Thalassiothrix frauenfeldii (Grunow) Castracane, Diatom. Challen-
ger Exped. (1886) 54-55, pi. 14, figs. 7, 8; Peragallo^ Diatom. Mar.
France (1897-1908) 321, pi. 131, fig. 15.
Cell linear, 0.22 to 0.29 millimeter in length, 0.004 to 0.005 in
breadth, forming large star-shaped clusters. Geographic dis-
tribution: Atlantic and Pacific Oceans, Sea of Japan, Eastern
and South China Seas. Localities 4 and 5.
ASTERIONELLA JAPONICA Cleve. Plate 9, fig. 9.
Gran, Nord. Plankton (1906) 118, fig. 160.
Asterionella glacialis Castracane, Diatom. Challenger Exped. (1886)
50, pi. 14, %. 1; Schroder, Phytopl. Warm. Meere (1906) 330-37;
Okamura, Littoral Diatoms Japan (1911) 11, pi. 13, fig. 56.
Length of valve, 0.11 to 0.205 millimeter; breadth on one end,
0.003, on the other end, 0.018. Geographic distribution: An-
tarctic, Atlantic, and Pacific Oceans; in Japanese waters known
from Shira-hama, Misaki, and Shima. Localities 2, 4, and 5.
GRAMMATOPHORA JAPONICA Grunow. Plate 10, fig. 13.
Grunow in V. Heurck, Synopsis (1880-85) pi. 103, fig. 18; Peragallo,
Diatom. Mar. France (1897-1908) 358, pi. 137, fig. 26.
Valve 0.052 to 0.06 millimeter in length, 0.028 to 0.03 in
breadth; stride 28 in 0.01 millimeter. Geographic distribution:
Pacific Ocean, Sea of Japan, Dairen. Localities 1 and 2.
112 The Philippine Journal of Science mi
GRAMMATOPHORA MARINA (Lyngby) Ktitzingr. Plate 2, fig. 15.
Kutzing, Bacillar. (1844) 128, pi. 17, fig. 24; W. Smith, Brit. Dia-
tom. (1853-56) 11, 42, pi. 42, fig. 314; Schutt, Bacillar. (1896) 106,
figs. 187, A-B; Peragallo, Diatom. Mar. France (1897-1908) 353,
pi. 137, figs. 6-8.
Cell 0.021 to 0.027 millimeter in length, 0.017 to 0.02 in
breadth; striae 22 in 0.01 millimeter. Geographic distribution:
Atlantic and Pacific Oceans, Mediterranean Sea, Sea of Japan,
Vladivostok, and Dairen.
SYNEDRA AURICULATA Karsten. Plate 10, fig. 2.
Karsten, Phytopl. Atlant. Ocean (1906) 173, pi. 30, figs. 18a, b.
Valve linear, straight, 0.8 to 1.2 millimeters in length, 0.006
in breadth. The ends inflated and shortly rounded, striae 15 in
0.01 millimeter. Geographic distribution : Atlantic Ocean. Lo-
cality 4.
SYNEDRA KOREANA sp. nov. Plate 9, figs. 3, 4, and 5.
Valve straight, lanceolate or linear, inflated in the middle part
forming a broad lanceolate pseudoraphe. Ends inflated, pro-
longed into rostrate apices. Length, 2.22 to 2.56 millimeters;
breadth in the middle, 0.011 to 0.013; striae, 10 in 0.01 milli-
meter. Localities 4 and 5.
NAVICULA PELLUCIDA Karsten. Plate 9, figs. 11 and 12.
Karsten, Phytopl. Antarkt. Meere (1905) 126, pi. 18, fig. 3.
Valve elliptic with elongated rounded ends. Length, 0.088 to
0.115 millimeter; breadth, 0.029 to 0.04; median line with the
terminal fissures indistinct. Axial area also indistinct; central
small. Striae 20 to 28 in 0.01 millimeter, obscure, thin. This
diatom should not be confused with the later-named N. pellucida
Cleve. Localities 2, 4, and 5.
NAVICULA (CISTULA) LORENZIANA Grunaw. Plate 8, fig. 16.
Grunow, Oster. Diatom. (1860) 547, pi. 3, fig. 3; Cleve, Synopsis Nav.
Diatom. (1894) 124; Peragallo, Diatom. Mar. France (1897-1908)
pi. 7, fig. 6; A. Schmidt, Atlas Diatom., pi. 212, figs. 51-56.
Valve broad, rectangular. Striae composed of elongated
puncta 15 to 18 in 0.01 millimeter. Rows of puncta 7 in 0.01
millimeter. Length of valve, 0.04 to 0.05 millimeter; breadth,
0.019 to 0.02. Geographic distribution: Littoral zone of Eng-
land, Balearic Islands, Adriatic, Campeche Bay, Port Jackson,
Yokohama. Locality 4.
46,1 Skvortzow: Diatoms of Korean Strait 113
NAVICULA (SCHIZONEMA) RAMOSISSIMA Agardh forma AMPLIA Grunow. Plate
10, fig. 3.
Schizonema amplius V. Heurck, Synopsis (1880-85) pi. 15, fig. 3;,
Peragallo, Diatom. Mar. France (1897-1908) pi. 12, fig. 9.
Valve linear-lanceolate with obtuse ends. Length, 0.064 to
0.07 millimeter; breadth, 0.012 to 0.017; striae, 12 in 0.01 milli-
meter. Geographic distribution: Atlantic and Pacific Oceans, a
benthonic species. Localities 1 and 2.
NAVICULA (SCHIZONEMA) MOLLIS W. Smith. Plate 8, figf. 15.
W. Smith, Brit. Diatom. (1853-56) 11, 77, pi. 58, fig. 365; V. Heurck,
Synopsis (1880-85) pi. 15, figs. 22, 23.
Schizonema albicans V. Heurck, Synopsis, pi. 15, fig. 20.
Schizonema torquatum V. Heurck, Synopsis, pi. 15, fig. 21.
Valve lanceolate, obtuse; length, 0.028 to 0.03 millimeter;
breadth, 0.008 to 0.009 ; striae, 18 in 0.01. Geographic distribu-
tion: Arctic America, Cape Sabine, Bahuslan, North Sea,
Adriatic. Localities 1 and 2.
NAVICULA KARIANA Gnmow var. MINOR Grunow forma JAPONICA forma nov.
Plate 8, fisr. 12.
Valve broadly lanceolate with rostrate ends. Length, 0.023
to 0.027 millimeter; breadth, 0.009; striae, 18 to 20 in 0.01 milli-
meter. The typical Navicula kariana Grunow x and var. minor
Grunow 2 and var. minor Grunow forma curta Cleve 3 are known
from Franz Josef Land, Sea of Kara, Cape Wankarema, Davis
Strait, and Cape Deschnew. Locality 1 .
PLEUROSIGMA LONGUM Cleve var. INFLATA Peragallo forma JAPONICA forma nov.
Plate 10, figr. 15.
Valve lanceolate, sigmoid, acute. Length, 0.136 to 0.15 milli-
meter; breadth, 0.017 to 0.02; strise, 17 in 0.01 millimeter.
Geographic distribution: The typical var. inflata is known from
the Mediterranean. Locality 4.
PLEUROSIGMA WANSBECKII Donkin. Plate 10, fig". 14.
Pleurosigma balticum var. wansbeckii Donkin in Peragallo, Diatom.
Mar. France (1890-91) 19, pi. 7, figs. 23, 24.
Valve linear. Length, 0.119 to 0.2 millimeter; breadth, 0.015
to 0.02. Geographic distribution: Sea of Kara and North Sea.
Locality 4.
1Arct. Diatom., 39, pi. 2, fig. 44.
'Aret. Diatom., 5; =N. frigida Grunow in Arct. Diatom., 39.
* Diatom. Exped. Vega (1883) 469, pi. 37, fig. 40.
259737 8
114 The Philippine Journal of Science 1931
GUINARDIA FLACCIDA (Castracane) Peragrallo. Plate 2, fig. 17.
Peragallo, Diatomiste 1 (1892) 107, pi. 13, figs. 3, 4.
Rhizosolenia ftaccida Castracane, Diatom. Challenger Exped. (1886)
74, pi. 29, fig. 4.
Henseniella baltica Schutt in De Toni, Sylloge Algarum (1894) 1425.
Guinardia baltica Schutt, Bacillar. (1896) 84, fig. 138; Okamura, Lit-
toral Diatoms Japan (1911) 4, pi. 9, fig. 15; Hustedt, Kieselalgen
(1929) 561-64, fig. 322.
Cell cylindrical, from 0.023 to 0.07 millimeter broad and two
to three times as long as broad, forming a long straight chain.
Chromatophores numerous, cross-shaped. Geographic distribu-
tion: Atlantic and Pacific Oceans, Mediterranean Sea, and Sea
of Japan. Localities 2, 4, and 5.
RHIZOSOLENIA ALATA Brightwell. Plate 10, figs. 9 and 10.
Brightwell, Quar. Journ. Micr. Sc. 6 (1858) 96, pi. 5, fig. 8; Pera-
gallo, Monogr. Rhizosol. (1892) 20, pi. 5, fig. 11; Diatom. Mar.
France (1897-1908) pi. 18, fig. 11; Hustedt in A. Schmidt, Atlas
Diatom. (1920) pi. 317, figs. 1-7; Hustedt, Kieselalgen (1929)
600; Okamura, Littoral Diatoms Japan (1911) 6, pi. 9, fig. 27.
Cell 0.17 to 0.4 millimeter in length and 0.011 to 0.013 in
breadth. Geographic distribution : Atlantic and Pacific Oceans ;
common in Japanese waters and known from Cape Goza, Shira-
hama, Province of Tosa, and Mikawa. Localities 2 and 4.
RHIZOSOLENIA ALATA Brightwell forma GRACILLIMA (Clcve) Grunow. Plate 10, figs.
11 and 12.
V. Heurck, Synopsis (1880^-85) pi. 79, fig. 8.
Rhizosolenia (alata var.) gracillima Clevb, Kongl. Sv. Vet.-Akad.
Handl. 18 (1881) 26, pi. 6, fig. 78; Hustedt in A. Schmidt, Atlas
Diatom. (1920) pi. 317, figs. £-10; Hustedt, Kieselalgen (1929)
601, fig. 345.
Cells about 0.006 to 0.0074 millimeter in breadth and 0.2 to
0.85 in length. Geographic distribution: Atlantic, Pacific, and
Indian Oceans, Mediterranean and Red Seas, Malay Archipelago,
New Zealand, Sea of Japan. Localities 2, 4, and 5.
RHIZOSOLENIA SETIGERA Brightwell. Plate 10, fig. 5.
Brightwell, Quar. Journ. Micr. Sc. 6 (1858) pi. 5, fig. 4;, V. Heurck,
Synopsis (1880-85) pi. 78, figs. 7, 8.
Rhizosolenia japonica Castracane, Diatom. Challenger Exped. (1886)
23, fig. 7; Peragallo, Monogr. Rhizosol. (1892) 17, pi. 4, figs. 12-16;
Diatom. Mar. France (1897-1908) 464, pi. 124, figs. 11-15; Oka-
mura, Littoral Diatoms Japan (1911) 5, pi. 9, fig. 22; Hustedt in
A. Schmidt, Atlas Diatom. (1920) pi. 320, figs. 6-8.
Valve linear, slightly siliceous, 0.75 to 0.9 millimeter in
length, 0.01 to 0.014 in breadth with structure hardly visible.
Spine long, thin; 0.12 to 0.14 millimeter in length. Geographic
46,1 Skvortzow: Diatoms of Korean Strait 115
distribution: Atlantic, Pacific, and Indian Oceans, Mediterra-
nean and Red Seas, Malay Archipelago, Sea of Japan. Localities
2, 4, and 5.
RHIZOSOLENIA ROBUSTA Norman. Plate 10, fig. 4.
Pritchard, Histor. Infusor. (1861) 866, pi. 8, fig. 42.
Rhizosolenia sigma Schutt, Pflanzenleb. d. Hochsee (1893) 22, fig.
2; Peragallo, Monogr. Rhizosol. (1892) 14, pi. 2, fig. 1; pi. 3,' figs."
1, 2; Diatom. Mar. France (1897-1908) pi. 123, figs. 1, 2; Karsten]
Indische Phytopl. (1907) 163, pi. 29, fig. 10; Hustedt in A. Schmidt,
Atlas Diatom. (1920) pi. 320, figs. 1-3; Kieselalgen (1929) 578-80,
fig. 330; Okamura, Littoral Diatoms Japan (1911) 4, pi. 9, fig. 18.
Cell robust, 0.13 to 0.22 millimeter in breadth, 0.5 to 0.7 in
length. The end is curved, contracted, pointed. Geographic
distribution : Indian and Pacific Oceans, Mediterranean and Red
Seas, Malay Archipelago, Sea of Japan. Localities 2, 4, and 5.
RHIZOSOLENIA HYALINA Ostenfeld. Plate 10, figs. 6, 7, and 8.
Ostenfeld and Schmidt, Plankt. Rode Hav Og Adenbugten (1901)
160-61, fig. 11; Hustedt in A. Schmidt, Atlas Diatom. (1920) pi.
319, figs. 11-13.
According to Cleve the original diagnosis of this diatom is as
follows :
Frustule very slightly siliceous (length 0.28 to 0.34 millimeter, width
0.028 to 0.032) ; structure hardly visible, squama te (4-5 squamae at the
same height) ; spine (0.032 to 0.04 millimeter long) very thin, slowly in-
crassated at the base; valve in a front view with a characteristic un-
dulation, of the contour and with a fissure, in which the spine of the
neighbour cell is fastened.
Geographic distribution: Red Sea, Japan (Binn-meer bei
Akashi, r.m. by Hustedt) . Locality 4.
NITZCHIELLA LONGISSIMA (Brebisson) Ralfs forma TYPICA V. Heurck. Plate 10,
fig. 1.
Nitzschia birostrata Smith, Brit. Diatom. 1 (1853-56) 42, pi. 14, fig.
117; V. Heurck, Synopsis (1880-85) pi. 70, figs. 1-2; Traite Diatom.
(1899) 404, pi. 17, figs. 568.
Valve with long horns, 0.5 to 0.7 millimeter in length. Geo-
graphic distribution: A cosmopolitan diatom known in many
places. Localities 1 and 2.
NITZCHIELLA LONGISSIMA (Brebisson) Ralfs forma PARVA V. Heurck. Plate 9.
fig. 7.
V. Heurck, Synopsis (1880-85) pi. 70, fig. 3; Traite Diatom. (1899)
404, pi. 17, fig. 568; Peragallo, Diatom. Mar. France (1897-1908)
293, pi. 124, figs. 16-18.
A delicate diatom, 0.29 to 0.3 millimeter in length and 0.009
in breadth. Geographic distribution: Atlantic and Pacific
116 The Philippine Journal of Science 1931
Oceans, in littoral zone in benthos and plankton. Localities 1,
2, and 5.
SURIRELLA GEMMA Ehrenberg var. OVATA var. nov. Plate 8, &g. 13.
Valve broad ovate. Length, 0.068 to 0.072 millimeter;
breadth, 0.037 to 0.04. Costse 3 in 0.01 millimeter. Typical
Surirella gemma 4 have more-elongated valves. Geographic dis-
tribution: Atlantic and Pacific Oceans in littoral zone and in
plankton. Localities 1 and 2.
BIBLIOGRAPHY
Bailey, I. W.
1854. Notes on new species and localities of microscopical organisms.
Washington.
Brightwell, Th.
1856. On the filamentous longhorned Diatomaceae. London.
1858. Remarks on the genus Rhizosolenia. London.
Castracane, A. F. de.
1886. Report on the Diatomaceae collected by H. M. S. Challenger
during the years 1873-76. Report of the Challenger Exped.
Botany 2 London.
1887. Contribuzione alia florula della Diatomee del Mediterranea.
Roma.
Cleve, P. T.
1873. Examination of diatoms found on the surface of the Sea of
Java. Stockholm.
1873. On diatoms from the Arctic Sea. Stockholm.
1878. Diatoms from the West-Indian Archipelago. Stockholm.
1881. On some new and little known diatoms. Stockholm.
1883. Diatoms collected during the expedition of the Vega. Stock-
holm.
1894. Planktonundersokningar, Cilioflagellater och Diatomaeeer. Stock-
holm.
1894-95. Synopsis of the naviculoid diatoms. Stockholm.
1896. Diatoms from Baffin's Bay and Davis Strait. Stockholm.
1897. A Treatise of the Phytoplankton of the Northern Atlantic and
its Tributaries. Upsala.
Cleve, P. T., and A. Grunow.
1880. Beitrage zur kenntnis der arktischen Diatomeen. Stockholm.
De Toni, J.
1894. Sylloge Algarum omnium hucusque cognitarum. Patavii.
Ehrenrerg, C. G.
1854. Mikrogeologie. Bd. 1-2. Leipzig.
4 Figured in A. Schmidt, Atlas Diatom, pi. 24, figs. 26-27, and in V. Heurck,
Synopsis pi. 74, pis. 1-3.
46,1 Skvortzow: Diatoms of Korean Strait H7
Gran, H. H.
1897. Bacillariaceae vom kleinen Karajakfjord. Bibliotheca Botanica,
Heft. 42. Stuttgart.
1897. Protophyta, Diatomaceae, Silicoflagellata and Cilioflagellata. aus
den Norske Nordavs-Expedition 1876-78, Heft. 24. Kristiania.
1900. Bemerkungen uber einige Planktondiatomeen. Nyt Magazin for
Naturvidenskaberne, Bd. 38. Kristiania.
1900. Diatomaceae from the Ice-floes and Plankton of the Arctic Ocean.
The Norwegian North Polar Expedition 4 (1893-96) No. 2.
Kristiania.
1904. Die Diatomees der Arktischen Meere. Jena.
1906. Nordisches Plankton. 19. Diatomen.
Gran, H. H., and K. Yendo.
1914. Japanese Diatoms. Christiania.
Greville, K. K.
1865. Description of new genera and species of diatoms from Hong
Kong. London.
1866. Description of new and rare diatoms. London.
1866. Descriptions of new and rare diatoms from the Tropics and
Southern Hemisphere. Edinburgh.
Gbunow, A.
1860-62. Die Osterreichischen Diatomaceen. Wien.
1863. Ueber einige neue und ungenugend bekannte Arten und Gattun-
gen von Diatomaceen. Wien.
1867. Reise seiner Majestat Navara um die Erde. Botanisch. Teil.
Bd. 1. Algen. Wien.
1884. Die Diatomeen von Franz Josephs-Land. Wien.
Hustedt, Fr.
1927-29. Die Kieselalgen, aus Dr. L. Rabenhorsts Kryptogamen-Flora.
Leipzig.
IKARI, J.
1926-28. On some Chaetoceras of Japan. 1-2. Bot. Mag., Tokyo.
1927. On Bacteriastrum of Japan. Bot. Mag., Tokyo.
Jorgensen, E.
1900. Protphyten und Protzoen im Plankton aus red norwegischen
Westkuste. Bergen.
Karsten, G.
1905-6. Das Phytoplakton des Antarktischen Meere nach dem Material
der deutschen Tiefsee-Expedition 1898-1899. Jena.
1906. Das Phytoplakton des Atlantischen Oceans nash dem Material
der deutschen Tiefsee-Expedition 1898-1899. Jena.
1907. Das Indische Phytoplakton. Jena.
Kutzing, F. T.
1844. Die Kieselschaligen Bacillarien oder Diatomeen. Nordhausen.
Lauder, H. S.
1864. Remarks on the marine Diatomaceae found at Hong Kong with
descriptions of new species. London.
1872. On new diatoms. London.
118 The Philippine Journal of Science um
Leuduger-Fortmorel, G.
1892. Diatomees de la Malaisie. Buitenzorg.
Mann, A.
1907. Report on the diatoms of the Albatross voyages in the Pacific
Ocean 1888-1904. Washington.
1925. Marine diatoms of the Philippine Islands. Washington.
Mereschkowsky, C.
1900-02. On Polynesian diatoms. Scripta Bot., St. Petersburg.
Okamura, K.
1907. Some Chaetoceras and Peragallia of Japan. Tokyo.
1911. Some littoral diatoms of Japan. Tokyo.
OSTENFELD, C. H.
1902. Marine Plankton diatoms, in J. Schmidt, Flora of Koh Chang,
Part 7. Copenhagen.
Ostenfeld, C. H., and J. Schmidt.
1901. Plankton fra del Rode Hav og Adenbugten.
Pavillard, J.
1911-13. Observations sur les Diatoms. Bull. Sc. Bot. France, T 50-60.
Paris.
1916. Recherches sur les Diatomees pelagique du gulfe du Lion.
1925. Report on the Danish Oceanographical Expeditions.
Peragajxo, H.
1888. Diatomees de Medoc. Toulouse.
1890-1. Monographie du genre Pleurosigma. Paris.
1892. Monographie du genre Rhizosolenia et de quelques genre voisin.
Paris.
Peragallo, H. and M.
1897-1908. Diatomees Marines de France. Paris.
Pritchard, A.
1861. A History of Infusoria. London.
Rattray, J. A.
1889. A revision of the genus Coscinodiscus Ehrenb. and of some
allied genera. Edinburgh.
Schmidt, A.
1873-1927. Atlas der Diatomaceenkunde, in verbindung den Herren
Schmidt, Grundler, Grunow, Janisch, Weissflog, Witt, Fricke,
and Fr. Hustedt, pis. 1-368.
Schutt, F.
1888. Ueber die Diatomaceengattung Chaetoceras. Leipzig.
1893. Das Pflanzenleben des Hochsee. Kiel und Leipzig.
1895. Arten von Chaetoceras und Peragallia. Berlin.
1896. Bacillariales. Leipzig.
Schroder, B.
1906. Beitrage zur kenntnis des Phytoplankton warmer Meere. Zu-
rich.
1911. Adriatisches Phytoplankton. Wien.
46,1 Skvortzow: Diatoms of Korean Strait 119
Skvortzow, B. W.
1929. Marine diatoms from Dairen, South Manchuria. Philip. Journ.
Sci. 38, No. 4, Manila.
1929. On some marine diatoms from Siberian shore of Japanese Sea.
Jap. Bot. Mag. 43, No. 506. Tokyo.
Smith, W.
1853-56. Synopsis of the British Diatomaceae. London.
Van Heurck, H.
1880-85. Synopsis des Diatomees de Belgique. Anvers.
1899. Traite des Diatomees. Anvers.
ILLUSTRATIONS
Plate l
Figs. 1 and 2. Coscinodiscus concinnus W. Smith.
Fig. 3. Coscinodiscus radiatus Ehrenb.
4. Coscinodiscus concinnus W, Smith, an abnormal valve.
Figs. 5 and 6. Coscinodiscus concinnus W. Smith. The middle parts of the
valves greatly enlarged.
Plate 2
Figs. 1 and 2. Stephanopyxis palmeriana (Grev.) Grun.
Fig. 3. Stephanopyxis palmeriana forma curta forma nov.
4. Stephanopyxis turris (Grev. and Arn.) Ralfs.
Figs. 5 and 6. Eucampia zodiacus Ehrenb.
7 and 8. Ditylium brightwellii (West) Grun.
Fig. 9. Eucampia biconcava (Cleve) Ostenf.
10. Thalassiosira hyalina (Grun.) Gran.
11. Lauderia borealis Gran.
12. Biddulphia pulchella Gray.
13. Schroederella delicatula (Per.) Pavil.
14. Leptocylindrus curvatus sp. nov.
15. Grammatophora marina (Lyngb.) Kiitz.
16. Biddulphia longicomis Grev.
17. Guinardia flaccida (Castr.) Per.
Plate 3
Fig. 1. Chaetoceras boreale Bail.
2. Chaetoceras javanicum Cleve.
3. Chaetoceras siamense Ostenf.
4. Chaetoceras lorenzianum Grun.
Plate 4
Fig. 1. Chaetoceras messanense Castr.
2. Chaetoceras affine Lauder.
3. Chaetoceras saltans Cleve.
Figs. 4 and 5. Chaetoceras peruvianum Brightw.
Plate 5
Fig. 1. Chaetoceras compressum Lauder.
2. Chaetoceras radians Schiitt.
3. Chaetoceras didymum Ehrenb. var. genuina Gran.
4. Chaetoceras protuberans Lauder.
5. Chaetoceras didymum Ehrenb. var. genuina Gran.
6. Chaetoceras didymum Ehrenb. var. anglica Gran.
7. Chaetoceras sociale Lauder.
121
122 The Philippine Journal of Science
Plate 6
Fig. 1. Chaetoceras dadayi Pavil.
2. Chaetoceras tortissimum Gran.
FlGS. 3 and 4. Chaetoceras decipiens Cleve.
Plate 7
Fig. 1. Chaetoceras ikari sp. nov.
2. Chaetoceras reichelti Hustedt.
Figs. 3, 4, and 5. Chaetoceras atlanticum Cleve.
Plate 8
Fig. 1. Bacteriastrum varians Lauder.
2. Bacteriastrum hyalinum Lauder.
3. Bacteriastrum varians Lauder.
4. Bacteriastrum minus Karsten.
Figs. 5, 6, and 7. Bacteriastrum varians Lauder.
Fig. 8. Bacteriastrum comosum Pavil. var. hispida (Castr.) Ikari.
9. Biddulphia sinensis Grev.
Figs. 10 and 11. Thalassiothrix nitzschioides Grun.
Fig. 12. Navicula kariana Grun. var. minor Grun. forma japonica forma
nov.
13. Surirella gemma Ehrenb. var. ovata var. nov.
14. Corethron pelagicum Brun.
15. Navicula (Schizonema) mollis W. Smith.
16. Navicula (Cistula) lorenziana Grun.
Plate 9
FlGS. 1 and 2. Thalassiothrix antarctica Schimper forma japonica forma
nov.
3, 4, and 5. Synedra koreana sp. nov.
Fig. 6. Thalassiothrix nitzschioides var. javanica Grun.
7. Nitzchiella longissima (Breb.) Ralfs forma parva V. Heurck.
8. Thalassiothrix frauenfeldii Grun.
9. Asterionella japonica Cleve.
10. Planktoniella sol (Wallich) Schiitt.
Figs. 11 and 12. Navicula pellucida Karsten.
Plate 10
Fig. 1. Nitzchiella longissima (Breb.) Ralfs forma typica V. Heurck.
2. Synedra auriculata Karsten.
3. Navicula (Schizonema) ramosissima Ag. forma amplia Grun.
4. Rhizosolenia robusta Norman.
5. Rhizosolenia setigera Brightw.
FlGS. 6, 7, and 8. Rhizosolenia hyalina Ostenf.
9 and 10. Rhizosolenia alata Brightw.
11 and 12. Rhizosolenia alata forma gracillima (Cleve) Grun.
Fig. 13. Grammatophora japonica Grun.
14. Pleurosigma wansbeckii Donk.
15. Pleurosigma longum Cleve var. inflata Perag. forma japonica
forma nov.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 46, No. 1.
;vr=$8
PLATE 1.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 46, No. 1.
PLATE 2.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 4fi, No. 1.
^A^i^wy^/^.^,,^
PLATE 3.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 4G, No. 1.
PLATE 4.
Skvobtzow: Diatoms of Korean Stbait.]
[Philip. Journ. Sci., 46, No. 1.
PLATE 5.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 46, No. 1.
PLATE 6.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 16, No. 1.
PLATE 7.
Skvortzow: Diatoms of Korean Strait.
[Philip. Journ. Scr., 4(5, No. 1.
PLATE 8.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 4G, No. 1.
PLATE 9.
Skvortzow: Diatoms of Korean Strait.]
[Philip. Journ. Sci., 46, No. 1.
PLATE 10.
THE HISTAMINE TEST AS AN AID IN THE DIAGNOSIS
OF EARLY LEPROSY
By Jose Rodriguez and Fidel C. Plantilla
Of the Philippine Health Service, Cebu, Cebu
It is generally agreed that one of the greatest needs in leprosy
work to-day is a reliable serological test that can be depended
upon to detect the disease in its earliest stages. Unfortunately,
in spite of claims of some to the contrary, such a test does not
yet exist. Until one has been elaborated and since in the "inci-
pient stage" the presence of Mycobacterium leprae cannot usu-
ally be demonstrated on ordinary methods of making the bac-
teriological examination, we have to depend almost entirely on
clinical methods such as the detection of the anaesthesia, palpa-
tion of thickened nerves and superficial glands, careful history-
taking, and examination of the external lesions as to appearance,
location, etc., in order to arrive at a diagnosis in this stage.
Naturally the accuracy of the diagnosis must depend to a great
extent on the experience of the physician making the diagnosis.
The introduction, therefore, of any clinical test that will tend
to minimize the influence of the personal equation should prove
of value.
We believe that we have found such a test in the so-called
"histamine test." When a dilute solution of histamine is pricked
into the normal skin, a reaction takes place in about twenty
seconds, starting with the appearance of a circular, sharply de-
fined, local reddening surrounding the prick, and measuring
when fully developed from 3 to 4 millimeters in diameter. This
is followed in another fifteen to thirty seconds by a flush or
"flare" that appears on the surrounding skin. It is of the
utmost importance to distinguish this flare from the local red
reaction. The flare is dark red or scarlet contrasting with the
brighter shade of the latter; it has diffused and often crenated
borders that may extend from 2 to 3 centimeters from the
center of the reaction. Soon after the appearance of the flare,
a discreet wheal forms at the site of the prick ; this is generally
at its maximum development in from three to five minutes, at
123
124 The Philippine Journal of Science 1931
which time it measures from 3 to 4 millimeters in diameter and
about 1 to 2 millimeters in height. The wheal usually occupies
the area originally covered by the local red reaction, although
in many cases the two do not coincide, the wheal being usually
smaller than the localized red area.
The full reaction of the normal skin to histamine, consisting
of the local redness or vasodilation, the flare, and the oedema or
wheal has been called by Lewis x the "triple response."
Lewis has demonstrated that the triple response is a char-
acteristic reaction of the normal skin following injury inflicted
by such agents as heavy stroking, pricking, scratching, freez-
ing, heating, electrical shocks, as well as by the introduction of
irritant substances such as acids, alkalies, mustard oil, ean-
tharidis, nettle sting, morphine, etc. Ultraviolet rays, ordinary
sunlight, X-ray and radium emanations, bacterial poisons, cer-
tain chemicals such as dichloraethylsulphide, etc., give rise to
more slowly developing reactions. He has also proven that the
local redness and the wheal or oedema are due to direct action
of the injury or irritant on the capillaries, while the flare is
produced by the dilatation of the arched arterioles and is reflex
in nature, being dependent upon the integrity of the cutaneous
nerves. The arteriolar dilatation is mediated through a purely
local nervous reflex and does not depend upon a spinal reflex arc.
This test has been tried by Lewis and his colleagues 2 on
anaesthetic skin to which the sensory nerves have been cut sur-
gically or interrupted by injection of anaesthetics. When the
interruption produced surgically or by anaesthesia is recent, the
reaction to the histamine test is complete in all its details, al-
though the skin has already been rendered anaesthetic; but if
sufficient time (six to fifteen days) is allowed for the nerve
to degenerate after surgical section or if the skin is anaesthetized
locally, the flare is lost. Under these circumstances, the local
red reaction and the oedema appear as in the normal reaction
of the skin.
Thus, the loss of the flare following a histamine test is a sign
of degeneration of the sensory nerves supplying the skin tested,
and possibly also of direct involvement of the nerve endings as in
local anaesthesia.
aThe Blood Vessels of the Human Skin and their Responses. Shaw &
Sons, Ltd., London (1927) 47.
2 Op. cit. 69-70.
46, i Rodriguez and Plantilla: Histamine Test in Leprosy 125
Histamine, or /Mminazolylethylamine, is described by Lewis
as "the amine produced when carbon dioxide is split from histi-
dine, a substance occurring naturally in the body and a protein
derivative." It was extracted by Barger and Dale 3 from the
intestinal mucosa, and was later thoroughly studied by Dale and
Laidlaw.4 The histamine test as applied on the skin was first
reported by Eppinger 5 and later elaborated by Sollman and Pil-
cher 6 and by Lewis and Grant.7
THE TEST
In most of our tests, we have used a 1 to 1,000 dilution of the
phosphate in normal salt solution. With stronger solutions a
larger flare is occasionally obtained, but the reactions are not
as constant as with the 1 to 1,000 solution.
A small drop of the solution is carefully placed within the
suspicious macule to be tested and another is dropped on normal
skin at least 2.5 centimeters from the border of the lesion for
control. With a sharp pin, a prick is made through the drop
into the skin underneath, taking care to exert just sufficient
pressure to drive the point through the epidermis without caus-
ing any bleeding. The histamine solution is wiped off im-
mediately, and the pricks are closely observed under good nat-
ural light.
The test is said to be negative when the complete response
is elicited and positive when the flare is absent.
There are some individuals on whom the normal reaction is
diminished; in a few, the flare is so faint as to be practically
absent. When the response is weak and the skin tested is on
an extremity, the flare may be brought out to its maximum
extent and intensity by previously congesting the extremity
with the help of a broad rubber band or the pneumatic cuff
of a blood-pressure apparatus.
Finally, it must be recognized that the reaction is harder to
elicit on the dark skin of a Filipino than on white skin.
8Journ. of Physiol. 41 (1910-11) 499-503.
4Journ. of Physiol. 41 (1910-11) 318-344; 43 (1911-1912) 182-195.
*Wein. med. Wochenschr. 43 (1913) 1414.
8 Journ. of Pharmacol, and Eper. Therap. 9 (1917) 309-340.
7 Vascular Reactions of the Skin to Injury. Part II, Heart 11 (1924) 209-
265.
126 The Philippine Journal of Science 1931
RESULT OF THE HISTAMINE TEST IN LEPROSY
In the pale macule. — The flush is always absent in the de-
pigmented macule of leprosy. When the histamine prick is
made just outside the border, a flare develops on the normal
skin but stops sharply at the border and does not extend into
the macule. When the prick is made just inside the border, the
flare is prevented from appearing even on the bordering normal
skin.
A word of caution must be given at this point. The flare
generally masks the local redness following the histamine test
on the normal skin. When the flare is abolished as in a leprotic
macule, the local redness becomes prominent and may be mis-
taken for the flare by the beginner. The area of local redness
is sharply localized, circular in shape, bright red or pink in
color, extending at the most 2 or 3 millimeters beyond the wheal,
and tends to become cyanotic before fading. On the other hand,
the flare is not definitely localized, the size is usually about 3
to 4 centimeters in diameter, irregular in shape, although it
tends to be oblong with its long axis along the length of the
member, and the color is dark red. On fading the flare becomes
speckled, but the color remains the same from beginning to end.
The wheal in the macule is usually of the same size as that
on the normal skin. Sometimes the oedema may be less; at
other times the wheal develops faster in the macule, reaching its
full development in two minutes, while the wheal on the con-
trol skin is at its height in three to five minutes. The ultimate
size, however, is almost the same.
The test has been applied on the macules of Tinea flava and
other types of pale-looking pityriases, on leucoderma, old scars,
fading psoriasis lesions, etc., which may be mistaken for the pale
macule of leprosy. In every case, the flare is present provided
the individual is not unsusceptible to histamine, in which case,
the flare is also diminished or absent on the normal skin.
In the reddish macule. — When the redness of the lesion is
marked, only the wheal may be elicited; but when the color
is not so striking, the local redness may be seen.
When hyperesthesia is present, as is usually the case when
the lesion is bacteriologically positive, the flare is not constant.
In a few macules the flare is present; in the majority of the
cases it is absent. If there is accompanying infiltration or
oedema so marked that the skin looks tense, glistening, and
bright red in color, the wheal is apt to be slight or absent.
46, i Rodriguez and Plantilla: Histamine Test in Leprosy 127
The histamine test was tried in cases of dermatitis from
various causes, active psoriasis lesions, tinea circinata and other
ringworm infections, fresh scars, and other lesions that may
simulate the red macule. When the inflammation in such lesions
is active and there is considerable redness, the wheal is gen-
erally diminished or even absent while the flare is present,
manifested by increased redness of the skin. It must be stated
that when the redness of the original lesion is at all bright, it is
next to impossible to distinguish the flare. When this is the
case, the best way to perform the test is to prick the histamine
solution just inside the border. In the nonleprotic lesion, the
flare appears on the adjacent portion of the skin outside the
border, whereas there is no such flare extending from the macule
in early leprosy.
SUMMARY
1. The histamine test has been found to be a fairly reliable
clinical test in differentiating the patches characteristic of the
early stages of leprosy from nonleprotic macules.
2. This test is "positive" (the flare is absent) in the large
majority of the bacteriologically negative leprotic macules tested.
3. The method of performing the test is described and its
limitations mentioned.
THE FLY EUTRIXOPSIS JAVANA TOWNSEND (DIP-
TERA, TACHINID^E), A PARASITE OF THE BEETLE
LEUCOPHOLIS IRRORATA IN OCCIDENTAL
NEGROS, PHILIPPINE ISLANDS
By A. W. Lopez
Chief Entomologist, Research Bureau, Philippine Sugar Association
On April 10, 1930, four maggots of the tachinid fly Entrixop-
sis javana Tns. were found in one specimen of the beetle Leuco-
pholis irrorata Chevr.,1 collected at Hacienda Candelaria, La
Carlota, Occidental Negros Province. The maggots pupated
April 11 and emerged April 20, a pupal period of nine days.
The determination of the flies was made by Dr. J. M. Aldrich,
of the United States National Museum, Washington, D. C.
The one specimen returned by Doctor Aldrich is in poor con-
dition, and it is impossible to give a description of it. However,
its length is 7 millimeters.
The publication Insecutor Inscitiae Menstruus 2 contains the
following description of one male collected at Pelaboean, Ratoe,
Java, by Bryant and Palmer.
Length 5.5 mm. wholly brownish-fulvous, including antennae and palpi;
tarsi darker, basal half or more of abdominal segments yellowish. Tegulae
tawny-whitish. Wings clear.
The United States Department of Agriculture3 reports that
E. javana was unwittingly introduced into the United States
from Sapporo, Japan, in 1922, with a shipment of material im-
ported to obtain Centeter cinerea, a tachinid parasite of the
Japanese beetle Popillia japonica. It is stated that the life cy-
cle of E. javana apparently corresponds closely to that of Cen-
teter, only one generation a year being produced.
1Coleoptera, Scarabaeidae, worst cane root-pest in the Philippine Islands.
2Nos. 10-12 6 (1918) 166.
3 Bull. 1429: 19-20, with fig. of E. javana.
259737 9 *29
COMPOSITION OF PHILIPPINE KAPOK-SEED OIL
By Aurelio 0. Cruz and Augustus P. West
Of the Bureau of Science, Manila
ONE PLATE
Kapok-seed oil is obtained from the seeds of the silk-cotton
tree (Ceiba pentandra Gaertner). Recently we determined the
composition of Philippine kapok-seed oil and our results showed
that this oil has a composition quite similar to that of American
cottonseed oil.
The silk-cotton tree is commonly known as kapok. It is a
tropical product and grows in tropical countries at such altitudes
as are free from frosts. In general, kapok is especially suited
to tropical lowlands. It is widely distributed in the Philippines.
The kapok tree is slender and usually has a height of about
15 meters or less. The branches are borne in horizontal whorls
that are very characteristic.
The kapok fruit is a capsule containing black seeds embedded
in fine silky hairs, or floss. The kapok fibers, or floss, surround-
ing the seeds are soft, elastic, and immune to moths. Kapok
floss has the property of being impermeable to moisture and is
also extremely buoyant. For this reason kapok is used exten-
sively for the manufacture of buoys, life belts, and life-saving
jackets. The chief use of kapok is for stuffing cushions, pillows,
mattresses, and similar articles. It is well adapted for this
purpose on account of its lightness, its springy or resilient
nature, and its nonhygroscopic and nonabsorbent characteristics.
Kapok floss is superior to most other flosses in resiliency and
consequently is more valuable for stuffing purposes. It has been
used considerably for making "down" quilts, which are about
as good as "eider down" quilts but much cheaper.
Kapok trees may be grown conveniently with other crops in
mixed plantation cultivation. The cultivation of crops under
kapok is quite practical because the few leaves and branches of
the kapok tree produce very little shade.
131
132 The Philippine Journal of Science 1931
Recently there have appeared several articles that give an
excellent account of the cultivation of kapok, the harvesting,
ginning, yields, insect pests, etc.1
Kapok trees begin to bear fruit in about four years, and when
seven years old they may yield about 500 pods per tree. The
yield naturally varies with the location and other factors. Un-
der favorable conditions much larger yields are obtained. The
number of pods required to produce a pound of clean floss is
said to average about 100.
Several years ago Philippine kapok appeared to be a very pros-
perous and promising industry. Like other products, however,
the value of kapok has decreased very considerably during the
recent financial depression. In 1927 the amount of kapok ex-
ported from the Philippines2 was 330,174 kilograms and the
value was 325,770 pesos. During 1929 there were exported
330,312 kilograms but the value was only 64,338 pesos. Prob-
ably when trade conditions are again adjusted kapok will return
to approximately normal values.
High-grade kapok-seed oil serves as an edible oil. The lower
grades are suitable for soap making and other purposes for
which low-grade cottonseed oil is employed. The oil cake left
after expression of the oil may be used for live-stock food or
fertilizer.
According to Lewkowitsch 3 kapok-seed oil is made in Holland
from seeds imported from Java. The oil gives color reactions
similar to those of cottonseed oil.
EXPERIMENTAL PROCEDURE
Philippine kapok pods, Ceiba pentandra, consist of about 51
per cent of husk and core, 32 per cent of seeds, and 17 per cent
of floss. One pod weighs about 32 grams and gives an average
of about 149 seeds, which weigh about 10 grams.
The Philippine kapok seeds used in this investigation were
kindly given to us by Dr. Manuel Roxas, director, Bureau of
Plant Industry. The seeds were ground in a mill after which
they were cold pressed to obtain the kapok oil. The oil was
purified by treating successively with 2 per cent Kieselguhr,
Suchar, and talcum powder. This treatment removes vegetable
1 Grist, D. H., Malayan Agr. Journ. 11 (1923) 3. Saleeby, M. M., The
Kapok Industry, Bull. Philip. B'ur. Agr. 26 (1922). Bull. Imp. Inst. 24 (1926)
18.
2 Annual Report, Insular Collector of Customs, Manila (1928 and 1930).
3 Chemical Technology and Analysis of Oils, Fats, and Waxes 2 (1922)
187.
4€,1
Cruz and West: Kapok-seed Oil
133
fibers and colloidal matter and produces a brilliantly clear yellow
oil with a slightly greenish tinge. The yield of oil calculated
on a moisture-free basis was found to be about 25 per cent.
The constants of this sample of Philippine kapok-seed oil are
given in Table 1.
Table I.— -Physical and chemical constants of Philippine kapok-seed oil
30o,
Specific gravity at — C.
4°
0.9109
1.4678
95.6
192.1
0.78
7.39
21.73
Refractive index at 30° C.
Iodine number (Hanus)
Saponification value
Unsaponifiable matter (per cent)
Acid value
Saturated acids, determined (per cent)
Unsaturated acids plus unsaponifiable matter, deter-
mined (per cent) 72.62
Saturated acids, corrected (per cent) 18.64
Unsaturated acids, corrected (per cent) 75.71
Iodine number of unsaturated acids plus unsaponi-
fiable matter 123.4
Iodine number of unsaponifiable matter 82.4
Iodine number of unsaturated acids (calculated) 123.9
The saturated and unsaturated acids that occur as glyce-
rides in Philippine kapok oil were separated by the lead-salt-
ether method 4 in accordance with the suggestions of Baughman
and Jamieson.5 The results are recorded in Table 2.
Table 2.— Separation of saturated acids from the unsaturated acids in
Philippine kapok-seed oil by the lead-salt-ether method.
Experiment No.
Oil used.
Unsaturat-
ed acids.
Saturated
acids.
Unsaturat-
ed • acids
(determin-
ed).
Saturated
acids (de-
termined) .
Unsaturat-
ed acids
(correct-
ed).
Saturated
acids (cor-
rected) .
1
9>
9.7005
11.5115
9.
7.0033
8.4079
9-
2.1209
2.4870
Per cent.
72.20
73.04
Per cent.
^ 21.86
• 21.60
Per cent.
75.25
76.17
Per cent.
18.81
18.47
2 __
Mean
72.62
21.73
75.71
18.64
a Iodine number (Hanus) of unsaturated acids plus unsaponifiable matter, 123.4.
b Iodine number (Hanus), 17.2.
c Iodine number (Hanus), 17.9.
The unsaturated acids separated from kapok oil by the lead-
salt-ether method were treated with bromine and converted into
their bromo-derivatives. No ether-insoluble hexabromide was
obtained, thus showing the absence of linolenic acid. The com-
4 Lewkowitsch, J., Chemical Technology and Analysis of Oils, Fats, and
Waxes 1 (1921) 556.
5 Cotton Oil Press 6 (1922) 41. Journ. Am. Chem. Soc. 42 (1920) 239S.
134
The Philippine Journal of Science
1931
position of the mixed unsaturated acids, which occur as gly-
cerides in kapok oil, was calculated from the iodine number of
the unsaturated acids. The results are recorded in Table 3.
There are also included the calculated percentages of glycerides
corresponding to these individual unsaturated acids.
Table 3. — Percentage composition of the unsaturated acids of kapok-seed
oil and the glycerides corresponding to these acids.
Acid.
Mixture of
unsaturat-
ed acids. *
Original oil.
Glycerides
in original
oil.
Linolic
Per cent.
37.02
62.98
Per cent.
28.03
47.68
Per cent.
29.29
49.83
Oleic
Total -
100.00
75.71
79.12
a Calculated iodine numbeT of the pure unsaturated acids was 123.9.
Saturated acids. — The saturated acids were separated from
Philippine kapok oil by the lead-salt-ether method and esterified
with methyl alcohol. The mixed acids were dissolved in methyl
alcohol and saturated with dry hydrogen chloride gas. The
mixture was then heated on a water bath (reflux) for fifteen
hours, after which it was treated with water and the ester layer
separated. The esters were dissolved in ether and the ethereal
solution washed with sodium carbonate solution and afterwards
with water. The ethereal solution was then dehydrated with
anhydrous sodium sulphate, filtered, and the ether removed by
distilling. The impure esters (83.97 grams), which were
yellow, were distilled under diminished pressure. A prelim-
inary distillation at about 3 millimeters pressure was made.
The esters (83.93 grams) were then redistilled at 3 millimeters
pressure. Data on the distillation of the esters are given in
Tables 4 and 5.
Table 4. — First distillation of the methyl esters of the* saturated acids;
pressure, 3 millimeters; 83.97 grams of esters distilled.
Fraction.
Temper-
ature.
Pressure.
Weight.
A
oc.
163-167
167-170
170-174
174-196
mm.
3
3
3
3
g-
19.43
19.98
17.07
22.34
5.10
B. - - -
C— — .
D— -
Residue - _
Total
83.92
46,1
Cruz and West: Kapok-seed OH
135
Table 5. — Second distillation of the methyl esters of the saturated acids $
pressure, S millimeters; 83 £2 grams of esters redistilled.
Fraction.
Temper-
ature.
Pressure.
Weight.
From first distillation.
Second
distillation.
A __ ---
1
2
3
4
5
Residue. __
163-167
167-169
169-175
175-192
192-222
mm.
3
3
3
3
3
9-
21.19
33.04
13.00
8.99
5.94
1.67
B and C
D -
Total
83.83
In Table 6, are given the analyses of fractions obtained in
the second distillation of methyl esters. From the data (Table
6), there were calculated the amounts of the individual acids
corresponding to the methyl esters contained in the various
fractions. The results are recorded in Table 7 and were cal-
culated in accordance with the methods outlined by Baughman
and Jamieson in their investigations of Hubbard squash-seed
oil 6 and also American cottonseed oil.7
In Table 8 is given the composition of the mixed saturated
acids and the glycerides in the original sample of kapok-seed oil
corresponding to these acids.
Table 6. — Analyses of fractions obtained in the second distillation of the
mixed methyl esters.
Fraction.
Iodine
number. »
Saponifica-
tion value.b
Mean
molecular
weight of
mixed es-
ters.
Composition of mixed
esters.
Mean
molecular
weight of
saturated
esters.
267.8
271.9
272.9
285.9
308.2
Saturated
Unsaturat-
ed.
1 ___
5.93
9.73
21.75
37.39
39.99
208.5
204.9
202.7
194.2
184.8
269.1
273.8
276.8
288.9
303.6
Per cent.
94.95
91.72
81.49
68.18
65.97
Per cent.
5.05
8.28
18.51
31.82
34.03
2
3
4 _-
5
» Calculated iodine number of unsaturated methyl esters was 117.5.
* Calculated saponification value of unsaturated methyl esters was 190.0.
•Journ. Am. Chem. Soc. 42 (1920) 156.
'Journ. Am. Chem. Soc. 42 (1920) 1197.
136 The Philippine Journal of Science 1931
Table 7. — Saturated acids corresponding to methyl esters in each fraction.
Fraction.
Acid.
Myristic.
Palmitic.
Stearic.
Arachidic.
1
Per cent.
7.96
9-
1.69
Per cent.
82.02
82.00
70.09
28.64
0.
17.38
27.09
9.11
2.58
Per cent.
ff.
Per cent.
9-
2
4.99
7.21
36.20
40.87
1.65
0.94
3.25
2.43
3
4
~-
5
22.10
1.31
1.59
Total__
1.69
56.16
8.27
2.90
a Residue assumed to be methyl arachidate.
Table 8. — Saturated acids.
Acid.
Mixture of saturated acids.
Glycerides
in original
oil.
Weight.
Composi-
tion.
Proportion
in original
oil.
9-
1.69
56.16
8.27
2.90
Per cent.
2.45
81.37
11.98
4.20
Per cent.
0.46
15.17
2.23
0.78
Per cent.
0.49
15.91
2.33
0.81
Total
69.02
100.00
18.64
19.54
The composition of Philippine kapok-seed oil is given in Table
9. There is also included for comparison the analysis of Amer-
ican cottonseed oid.
Table 9. — Composition of Philippine kapok-seed oil compared with
American cottonseed oil.
Constituent.
Glycerides of:
Unsaturated acids-
Oleic
Linolic
Saturated acids —
Myristic
Palmitic
Stearic
Arachidic
Unsaponiflable matter __
Total.
Philippine
kapok-seed
oil.
Per cent.
49.8
29.3
0.5
15.9
2.3
0.8
0.8
99.4
American
cottonseed
oil.*
Per cent.
35.2
41.7
0.3
20.0
2.0
0.6
99.8
a Composition determined by J. S. Jamieson and W.
Soc. 42 (1920) 1197.
F. Baughman, Journ. Am. Chem.
46,1 Cruz and West: Kapok-seed Oil 137
The determined iodine number of Philippine kapok-seed oil
was found to be 95.6 and the determined saponification value
192.1. The calculated iodine number is 93.8 and the saponifica-
tion value 191.3. The iodine and saponification values cal-
culated from the composition of the oil agree very closely with
the determined values.
SUMMARY
Kapok floss is an excellent material for stuffing cushions, pil-
lows, mattresses, buoys, life-saving jackets, and similar articles.
It is well adapted for this purpose on account of its lightness,
its springy or resilient nature, and its nonhygroscopic and non-
absorbent characters.
Kapok can be grown in the Philippines conveniently with
other crops in mixed plantation cultivation.
The composition of Philippine kapok-seed oil has been deter-
mined, and the results (Table 9) indicate that the Philippine oil
has a composition very similar to that of American cottonseed
oil.
The percentage of linolic and palmitic glycerides is slightly
higher in the cottonseed oil than in the kapok oil. The kapok-
seed oil has a higher percentage of oleic glyceride than the
cottonseed oil, while the percentage of the other glycerides is
about the same.
Since Philippine kapok floss is superior to most other flosses
and kapok seeds yield an oil of high quality and of about the
same composition as American cottonseed oil, it would seem
that there are promising prospects for the development of kapok
cultivation in the Philippines under normal trade conditions.
ILLUSTRATION
Plate 1, Philippine kapok trees and seed pods.
139
Cruz and West; Kai*ok-skki» ou..\
| Philip. Journ. Sri.. 46, No. 1.
^.-,
*' -i\ . *
rs"
v ' T^W"t* •
M^M ' **- ^
>LATE 1. PHILIPPINE KAPOK TREES AND SEED PODS.
THE SKELETON OF THE TIMARAU
By Manuel D. Sumulong
Of the College of Veterinary Science
University of the Philippines
Los Banos, Laguna
THREE PLATES AND FOUR TEXT FIGURES
This paper records some observations on the characteristic
features of the skeleton of the full-grown timarau (also spelled
timerau and tamarao), a wild species of the family Bovidse,
confined to Mindoro Island, and the largest indigenous mammal
of the Philippines. A careful search of the literature on hand
has shown nothing about the internal features of this animal.
With the hope of partially filling this gap in our knowledge the
present study was undertaken.
In the following account an attempt will be made to point
out where the skeleton of the animal in question differs from
those of the carabao and the cow, making the descriptions of
the individual bones comparative, where comparison is possible.
Concerning the habitat, size, and external features of this
animal, Steere (1888) states in part:
I have been in the interior of the little-known island of Mindoro, and
have had the satisfaction of procuring specimens of a strange animal
there, which, though generally talked of throughout the Philippines, is
little known to scientific men. This is the "Tamaron". From the native
reports I could make out nothing, but that it was a large fierce beast with
sharp horns, which attacked all who came near it . . .
In Mindoro I procured three full-grown individuals (two males and one
female) of the "Tamaron," and have preserved the skins and skeletons . . .
General color of the skin and hair black, hair short and rather fine. A
grayish-white stripe running from near the inner corner of the eye to-
wards the base of the horn (this stripe three inches long by one inch
wide), a grayish-white spot above each hoof on all feet, a grayish-white
patch on inner side of lower foreleg; skin and hair of groin white;
bare skin of nose and lips black; horns and hoofs black; tips of horns
pointed and polished; horns triangular, with a tendency in the bulls to-
wards thickening and flattening at the base; lower part of the horns
with deep irregular pits; several of the last vertebrae of the tail aborted.
Size of No. 1 : An old bull : length from point of nose to tip of tail eight
feet one inch; length of tail one foot five inches; length of tassel of hairs
at end of tail two and a half inches; height at shoulder three feet six
141
142 The Philippine Journal of Science 1931
inches; from breast-bone to sole of fore foot one foot eight inches; length
of horn one foot two inches; circumference of horns at base thirteen in-
ches; horns distance apart at base one and a half inches, at points ten
inches; length of head, before skinning, one foot four inches.
Montellano (1929) describes the timarau as follows:
The tamaraw is much like the domesticated carabao, except in size,
shape, and size of horns, and conformation of the body. The tamaraw
is smaller. The horns are rather short and straight, point vertically up-
ward, and gradually taper to a sharp point admirably adapted for fighting.
The body is lighter and shallower, and is better adapted to rapid move-
ment than is that of the domesticated carabao.
These animals, however, are not the ancestors of the wild carabaos
found elsewhere in the Philippines, and in Borneo and other neighboring
islands, and in Southern Asia.
According to Sclater (1888), Steere proposed to call this
species Anoa mindorensis, because of its very close resem-
blance to the Anoa of Celebes ; but Bubalus mindorensis, as pro-
posed by Heude (1888), seems to have been finally accepted.
Meyer (1878) is of the opinion that the timarau is entirely
different from the Anoa of Celebes. Bartlett as well as Gray
(1878) believes that it is but a "small variety of the common
Manila or water Buffalo." As reported by "Peres de la Cam-
pagnie de Jesus (1888) ," it is not at all a type of ordinary buffalo.
Their report asserts that a buffalo that has escaped from its
owner and has become wild for a long time will never produce
a timarau. This agrees with the observation of Steere that the
timarau is distinctly different from the so-called "carabao ce-
maron," a wild carabao found in the Philippines, especially
in Luzon.
Mention of the timarau as a source of food supply has been
made by Miller (1912) in his study of the Mangyans of Min-
doro.
In the preparation of this paper the previous work of the
writer (1926) on the skeleton of the carabao {Bubalus bubalis),
the Filipino beast of burden, was freely consulted.
MATERIAL
The data presented here were obtained from a thorough
study of the mounted skeleton of an adult timarau in the ana-
tomical museum of the College of Veterinary Science, Univer-
sity of the Philippines, and of another in the museum of the
University of Santo Tomas, Manila. So far as the writer is
aware these are the only mounted skeletons of this animal in
the Philippines. The skeleton at Santo Tomas University is
46, i Sumulong: Skeleton of the Timarau 143
incomplete, the mandible as well as some of the small bones
of the limbs being lacking, and no data could be obtained as
to the history of the animal from which it was prepared. The
specimen at the College of Veterinary Science is a complete ar-
ticulated skeleton of an adult timarau which, so far as the
writer could recall, was presented to the College in 1916 by
an American gentleman with the request that it be sacrificed.
OSTEOLOGY
The following are the bones of the various regions of the
skeleton of the timarau:
THE AXIAL SKEI*BTON
A. The Skull.
1. Bones of the cranium.
a. Single bones.
1. Occipital.
2. Sphenoid.
3. Ethmoid.
b. Paired bones.
1. Interparietal.
2. Parietal.
3. Frontal.
4. Temporal.
2. Bones of the face.
a. Single bones.
1. Vomer.
2. Hyoid.
3. Mandible.
b. Paired bones.
1. Maxilla.
2. Premaxilla.
3. Nasal.
4. Malar.
5. Lacrimal.
6. Pterygoid.
7. Palatine.
8. Dorsal turbinate.
9. Ventral turbinate.
B. The Trunk.
1. The vertebral column.
a. Cervical vertebrae, 7.
6. Thoracic vertebrae, 13.
c. Lumbar vertebrae, 6.
d. Sacral vertebrae, 5.
e. Coccygeal vertebrae, 15.
2. The thorax.
a. Ribs (both sides), 26.
b. Sternum (7 sternebrae), 1.
144 The Philippine Journal of Science 193 1
THE APPENDICULAR SKELETON
A. Bones of the Thoracic, or Pectoral, Limb.
a. Shoulder.
1. Scapula (both sides), 2.
b. Arm.
1. Humerus (both sides), 2.
c. Forearm.
1. Radius (both sides), 2.
2. Ulna (both sides), 2.
d. Manus.
1. Carpus (both sides), 12.
2. Metacarpus (both sides), 2.
3. Digits.
a. Phalanges (both sides), 20.
6. Sesamoids (both sides), 12.
B. Bones of the Pelvic Limb.
a. Pelvic girdle.
1. Os coxae (both sides), 2.
b. Thigh.
1. Femur (both sides), 2.
c. Leg.
1. Tibia (both sides), 2.
2. Fibula (both sides), 2.
3. Patella (both sides), 2.
d. Pes, or hind foot.
1. Tarsus (both sides), 10.
2. Metatarsus (both sides), 4.
3. Digits.
a. Phalanges (both sides), 20.
b. Sesamoids (both sides), 12.
In the preceding enumeration mandible, hyoid, and sternum
are regarded as single bones, and the os coxae is not divided
into its original parts — ilium, ischium, and pubis. The visceral
or splanchnic bones as well as the auditory ossicles are not in-
cluded.
For the purpose of giving an idea of the difference in size
between the skeleton of the timarau and that of the carabao,
measurements of the various parts of the mounted skeleton
of the timarau of the College of Veterinary Science and that of
a medium-sized adult carabao were taken. The length or height
of the flat and long bones of both the thoracic and pelvic limbs
were likewise determined. The results are given in Tables
1 and 2.
46,1
Sumulong: Skeleton of the Timarau
145
Table 1. — Showing the measurements of the various segments or regions of
the articulated skeleton of a timarau and of a medium-sized carabao.
[Measurements
in centimeters.]
Region.
Length.
Height.
Width.
Depth.
Circum-
ference.
Excess
in favor
of
cara-
bao.
Cara-
bao.
Tima-
rau.
Cara-
bao.
Ti-
ma-
rau.
Cara-
bao.
Tima-
rau.
Ca-
ra-
bao.
Tima-
rau.
Ca-
ra-
bao.
Ti-
ma-
rau.
Vertebral column
Tail_.-_
175.0
__68.0
123.0
35.0
35.5
52.0
33.0
15.0
5.5
8.5
18.5
13.0
10.0
51.5
33.0
33.0
Skull
50.5
20.0
14.5
Horn core... _J
Between bases of horn
cores
30
21.5
34.0
15.5
28
15
17.5
73.0
7.5
21.5
Between points of horn
Thoracic limb
115
118
82
85
Pelvic limb
Table 2. — Showing the height or length of the individual long and flat
bones of the appendicular skeleton of a timarau and of a medium-sized
carabao.
[Measurements in centimeters.]
Thoracic limb.
Pelvic limb.
Bone.
Cara-
bao.
Tima-
rau.
Excess
in favor
of cara-
bao.
Bone.
Cara-
bao.
Tima-
rau.
Excess
in favor
of cara-
bao.
Scapula
34.0
27.5
29.5
37.0
18.0
5.5
3.5
6.5
22.5
20.0
21.0
29.5
12.0
4.5
3.0
4.0
11.5
7.5
8.5
7.5
6.0
1.0
0.5
2.5
Os coxa?
48.0
38.0
32.0
20.0
6.5
4.5
7.5
32.0
28.0
24.5
14.5
5.0
3.5
4.5
16.0
10.0
7.5
5.5
1.0
1.0
3.0
Humerus
Femur
Radius _
Tibia
Ulna ___
Large metatarsal
First phalanx
Large metacarpal
First phalanx
Second phalanx _
Third phalanx
Second phalanx
Third phalanx
The distance between the level of the foramen magnum and
that of the posterior aperture of the sacral canal constitutes the
length of the vertebral column indicated in the table. The
length of the skull here was measured from the nuchal crest
to the central incisor teeth; the width refers to the broadest
part of its frontal surface, measuring along an imaginary line
259737-
-10
146 The Philippine Journal of Science wai
connecting the two supraorbital foramina. The depth refers to
the broadest part of its lateral surface including the mandible,
and w#s determined by measuring the distance between the
angle of the mandible and the level of the most prominent part
of the frontal region just in front of the base of the horn core.
The height of the anterior limb constitutes the distance, in a
straight line, between the highest point of the anterior or cer-
vical angle of the scapula and the ground plane, whereas that
of the posterior limb, is the distance between the highest point
of the tuber coxae and the ground plane.
THE SKULL
BONES OF THE CRANIUM
Occipital. — The occipital bone is very much less extensive than
that of the carabao or ox. The external surface of the squamous
and lateral parts, when taken as a whole, instead of being flat-
tened as in the carabao, is convex transversely. The nuchal
crest is markedly better developed, but the external occipital
protuberance is only represented by a rather faint elevation,
which is flanked on either side by a depression. The median
occipital crest is only represented here by a low ridge and does
not reach the upper border of the foramen magnum, fading out
halfway between its margin and the external occipital protube-
rance. It terminates into a rather deep depression bounded on
either side by a rounded muscular eminence formed by the fu-
sion of the squamous and lateral parts. The foramen magnum
is comparatively smaller, and its roof is perforated by three
small foramina located a short distance from its margin. The
paramastoid processes are short, being about one-half the length
of those of the carabao. The basilar part is likewise relatively
shorter and does not form with the body of the sphenoid pro-
minent ventral tubercles. The edge dividing the articular sur-
face of the condyle into an upper and a lower facet is better
defined. Except in size the hypoglossal and mastoid foramina
present no striking features.
Sphenoid. — The body of the sphenoid is narrow and short and
the temporal and orbital wings are less extensive. As in the
carabao there is a deep pituitary fossa and a very well-devel-
oped dorsum sellae. The foramen orbito-rotundum as well as
the foramen ovale presents no special features other than its
small size. The pterygoid crest is not well developed.
Ethmoid. — The ethmoid and its cells are well developed, dif-
fering only from those of the carabao in size. No attempt was
46,1
Sumulong: Skeleton of the Timarau
147
4 5'
Fig. 1. Lateral view of the skull of the timarau. a, Part of the frontal bone; b, lacrimal;
c, malar ; d, nasal ; e, maxilla ; /, premaxilla ; g, temporal ; h, occipital ; i, hyoid ; j, man-
dible; 1, horn core; 2, lacrimal fossa; 3, infraorbital foramen; 4. mental foramen; 5,
maxillary tuberosity ; 6, temporal fossa ; 7, coronoid process of the mandible ; 8, temporal
crest; 9, mastoid process; 10, external acoustic meatus; 11, paramastoid process; 12,
zygomatic arch ; IS, lacrimal bulla.
made to discover an air sinus in its perpendicular plate, which
is sometimes observed in carabao.
Interparietal. — The interparietal is completely fused behind
with the supraoecipital. Its external surface is smooth and flat
instead of convex as in the carabao; the cranial aspect is like
that of the same bone in the carabao or cow, carrying no distinct
tentorium osseum.
Parietal. — The external parietal crest is curved and better
defined than that of the carabao. This crest distinctly divides
the parietal bone into an upper horizontal part and a vertical
lower part. From the union of the horizontal parts of the two
parietal bones results a central plate whose anterior part is
triangular and concave. This is the only part of the parietals
that is visible when the skull is viewed directly from the front.
The posterior part that lies behind the line joining the bases
of the horn cores of the frontal bones is more or less quadri-
lateral in outline, presenting a comparatively smooth and slight-
ly convex outer surface which looks directly upward. The lower
vertical part that forms part of the medial wall of the temporal
fossa is slightly convex from the front backward, and it is
not concave from above downward as in the case of the carabao
and cow. Its anterior border is nearer to the frontal crest
than in the carabao.
148 The Philippine Journal of Science 1931
Frontal — The frontal bone is relatively narrower transversely
than in the carabao. Externally the nasofrontal part is more
concave in front, but nearly flat behind. There is no indication
of the frontal eminence at the junction of its posterior
border and the parietal bone. The horn cores are relatively
smaller, shorter, and less curved than in the carabao. They
are more or less three sided and taper to a blunt point. They
are directed almost straight backward, turning toward each
other moderately at the points ; they also run a little downward
bringing the ends to lie in the line of the orbit. The supraor-
bital foramen is relatively smaller and is placed higher. The
groove leading from it is narrower but deeper. The supraor-
bital process is weaker but relatively longer than in the carabao
or cow. The orbital part as well as the temporal part of the
bone is less concave and extensive.
Temporal. — The temporal bone of the timarau is character-
ized by the following features: The temporal crest is poorly
developed, and the zygomatic process is not as strong as in the
carabao. The external aspect of the part of the squamous tem-
poral that concurs with the parietal in the formation of the
medial wall of the temporal fossa is moderately convex, instead
of being concave as in the carabao or cow. The postglynoid
process is very poorly developed. The posterior process forms
a distinct muscular eminence behind the external acoustic pro-
cess. Aside from the difference in size, the muscular process,
the bulla ossea and the acoustic process present no other features
of interest.
BONES OF THE PACE
The bones of the face, aside from the difference in size,
present only a few important special features as compared with
those of the carabao.
Maxilla. — The facial tuberosity of the maxilla is only repre-
sented by a slightly elevated rough area, placed about an inch
above the alveolus of the third premolar tooth ; from it extends
backward and upward an ill-defined ridge which gradually
fades out and terminates at the junction of the maxilla and
the malar bone. The infraorbital foramen is relatively small
and is located just in front of the level of the alveolus for the
first premolar tooth. The maxillary tuberosity is very poorly
developed and very much compressed laterally ; it bears a short
blunt-pointed process that projects upward and backward. As
in the carabao this bone does not form any defect in its nasal
46, i Sumulong: Skeleton of the Timarwu 149
wall for it directly articulates with the nasal bone, and the
interval they form is completely occupied by the posterior ex-
tremity of the nasal process of the premaxilla. The anterior
part of the palatine process is narrow and is deeply concave
transversely. The maxillary foramen is slitlike and small.
Premaxilla. — The body of the premaxilla is relatively thin
and small, otherwise it resembles that of the carabao ; the fora-
men incisivum is represented by a notch. The palatine process
is practically as long as in the carabao and its posterior end
is overlapped by the anterior end of the vomer; it is deeply
grooved in the nasal surface for the reception of the ventral
edge of the vomer. The palatine fissure is narrow. The nasal
process is well developed and more or less prismatic; its poste-
rior end completely occupies the interval of the nasal and max-
illary bones.
Palatine. — The palatine bone closely resembles that of the
carabao.
Nasal. — The nasal bone, except in size, does not present many
important differential features that will attract attention. The
lower end of this bone is divided by a notch into an outer and
an inner process ; the latter is the smaller, instead of being the
larger, as is the case in the carabao.
Lacrimal. — The bulla of the lacrimal bone is proportionately
larger. In other respects this bone resembles that of the ca-
rabao.
Malar. — The facial part of the malar bone bears a less dis-
tinct ridge as compared with that of the carabao. This ridge
is apparently the continuation of the ill-defined crest of the
maxilla. The region behind the crest is less concave dorso-
ventrally. The upper extremity of the bone is bifurcate, the
upper branch being relatively shorter and weaker than in that
of the carabao or cow. The bone does not curve very much
laterally.
Aside from the difference in size the pterygoid bone does not
materially differ from the same bone in the carabao.
Vomer, hyoid, and turbinate. — The vomer, hyoid, and turbi-
nate bones resemble those of the carabao practically in all
respects.
Mandible. — The mandible likewise resembles very closely that
of the carabao in general form. The outer aspect of the per-
pendicular part of the ramus, however, is comparatively smooth,
presenting very few and less salient muscular ridges.
150
The Philippine Journal of Science
1931
THE SKULL AS A WHOLE
The skull of the timarau resembles in most respects that of
the carabao. The upper half of the frontal surface is relatively
narrower than in the latter animal, and it presents a slightly-
depressed central area. The
roof of the cranium is almost
flat. There is no indication at
all of the so-called median "fron-
tal eminence." The supraor-
bital foramen is placed higher
and the horn cores are more or
less three^sided and compara-
tively smaller and shorter; they
run almost straight backward
and a little downward. The zy-
gomatic arches and supraor-
bital processes do not curve out-
ward as much as in the carabao.
The following are the most
salient differential features of
the lateral surface: The facial
tuberosity is only represented by
a slightly raised rough area,
and the curved crest that ex-
tends from it is ill-defined and
incomplete. The temporal fossa
encroaches more on the poste-
rior surface ; it is relatively shal-
lower than in the carabao and
its medial wall is moderately
convex from before backward.
The external parietal crest,
which limits the fossa behind, is
better defined. At the junction
of the anterior extremity of the
body of the maxilla and the na-
sal process of the premaxilla is
a thin triangular plate of bone
projecting downward and outward.
The cranial part of the basal surface is relatively narrower
than in the carabao, and the tubercles in front of the occipital
condyles as well as the ventral tubercles at the junction of the
occipital and sphenoid bones are rather poorly developed. The
Fig. 2. Frontal view of the skull of the
timarau. a, Fused interparietal and su-
praoccipital ; 6, fused dorsal or horizon-
tal parts of the parietals ; c, frontal
bone ; d, lacrimal ; e, malar ; /, nasal ; g,
maxilla; h, premaxilla; 1, horn core (pro-
cessus corn us) ; 2, external parietal
crest ; S, supraorbital foramen ; -4, supra-
orbital groove; 5, orbit; 6, maxillary
tuberosity ; 7, nasal process of prema-
xilla; 8, palatine process of premaxilla;
9, palatine fissure; 10, palatine (notch)
cleft.
46, i Sumulong: Skeleton of the Timarcw 151
posterior nares are completely divided medially by the vomer.
The anterior palatine foramina are also found at the junction
of the horizontal part of the palatine bone and the palatine
process of the maxilla. The triangular plate of bone result-
ing from the union of the anterior extremity of the body of the
maxilla and the nasal process of the premaxilla is also visible
in this surface.
The posterior surface (nuchal surface) is distinctly divided
by a better-developed nuchal crest into an upper and a lower
area. The upper area is formed by the frontals, interparietals
and supraoccipital ; it is more or less quadrilateral in outline
and less extensive than the lower one ; it is smooth and slightly
convex from side to side ; and it is separated from the temporal
fossa by the parietal crests. The area below the nuchal crest is
rough and wide transversely below; the external occipital pro-
tuberance consists only of a small rough elevation flanked on
either side by a depression. Extending from this elevation is
a faint ridge that terminates below into a depression bounded
on either side by a rounded muscular eminence formed at the
junction of the squamous and lateral parts of the occipital bone.
Other features of this surface resemble very closely those of
the carabao.
The cranial cavity as well as the nasal cavity, aside from
the difference in size, is practically the same as in the carabao.
Mention may be made here that no attempt was made to open
and study the paranasal sinuses because we did not feel justified
in destroying the only mounted skeleton of the timarau in the
College.
THE VERTEBRAL COLUMN
The number of bones observed in each region of the verte-
bral column of the timarau is indicated in the following for-
mula, each region being denoted by its initial letter; C7 T13
L6 S5 Cy15. As to the number of bones, the sacral region of
the vertebral column of this animal differs from that of the
carabao, being made of only four segments or vertebrae, and
in the case of the ox the difference lies in the coccygeal region,
the number of coccygeal vertebrae in the latter animal varying
from 18 to 20.
Cervical vertebrae. — Except in respect to size these bones
resemble very closely those of the carabao. With the atlas and
axis, however, the following points are noteworthy : The tubero-
sity of the dorsal arch of the atlas is relatively better developed
than in the carabao, resembling very closely that of the ox.
152 The Philippine Journal of Science 1931
The wings are relatively thinner and less horizontal, and the
posterior border of the dorsal arch is deeply notched. The
fossa atlantis is shallower. The spinous process of the axis
is comparatively weaker and its free border is less tuberous.
The intervertebral foramen is placed farther behind, and the
foramen transversarium is relatively small. The transverse pro-
cesses are directed downward, outward, and backward, instead
of being horizontal as in the case of the carabao.
Thoracic vertebrae. — As compared with the same bones in
the carabao, these vertebrae do not present any striking dif-
ferential features, save that they are smaller and less voluminous
and that the free ends of the spinous processes are less tuber-
culate. Besides, both the anterior and posterior edges of the
spinous processes are straighter and more regular.
Lumbar vertebrae. — Aside from the difference in size these
vertebrae do not materially differ from those of the carabao.
The mammillary processes of these bones, however, are less
prominent and not as tuberous as in the latter animal; and
the transverse processes are relatively weaker and their edges
are more regular.
Sacrum. — This bone is relatively longer but less voluminous
than in the carabao; it is made of five segments or vertebrae
as in the ox. It is less arched. The spinous processes are
lower, and only those of the second, third, fourth, and fifth
vertebrae are completely fused together. The lateral borders
are not very thin, sharp, and irregular. The pelvic surface
is less concave in both directions, and the central groove is
hardly traceable.
Coccygeal vertebrae. — There are only fifteen coccygeal ver-
tebrae. A complete arch is present in the first seven bones,
which possess also transverse processes and distinct, though
nonfunctional, anterior articular processes. The arches as well
as the transverse and articular processes become more or less
rudimentary as they are traced backward. It may be remarked
here that the transverse processes of the first vertebra resemble
very closely those of the last segment of the sacrum both in
development and size, so that by casual observation it appears
to be a component of the sacrum which has not fused.
THE THORAX
As in the carabao the ribs of the timarau number thirteen
pairs — eight sternal and five asternals. They are proportion-
ately shorter, narrower, but more strongly curved than in the
46,1
Sumulong: Skeleton of the Timarcm
153
carabao. The necks are relatively
shorter, and the facets of the tuber-
cles are not deeply concave. The
borders are more regular.
The sternum consists also of seven
sternibrse and resembles that of the
carabao in general form. It is, how-
ever, relatively shorter and placed
less obliquely. The first sternebra is
not so bent upward, and the thorax
is more barrel-shaped than in the
carabao or ox.
THE BONES OF THE THORACIC LIMB
Scapula. — The scapula, of the tima-
rau resembles very closely that of the
carabao in general form, but in size
it is relatively smaller. The spinous
process is more sinuous and the tuber
spinse is less tuberous and poorly de-
veloped. The acromion is hardly re-
cognizable. The supraspinous and
inf raspinous fossae are shallower and
the anterior border is regular. The
tuber scapulae, as well as its coracoid
process, is less pronounced.
Humerus. — As compared with the
same bone in the carabao, the hum-
erus of the timarau presents the fol-
lowing features that are worth notic-
ing : It is shorter and less voluminous.
The musculospiral groove is deeper;
the deltoid tuberosity is more pro-
nounced; the teres tuberosity is ill-
defined; the curved line extending
from the deltoid tuberosity to the
neck is hardly distinguishable; and
the nutrient foramen is located about
the middle of the medial surface.
Radius and ulna. — Except in point
of size and the poorly developed ra-
dial tuberosity, the radius and ulna
are almost identical with those of
Fig. 3. Lateral aspect of the arti-
culated bones of the thoracic limb
of the timarau. a, Scapula; b,
radius ; c, radial carpal ; d, fused
second and third carpals ; e,
fourth carpal; f, large metacar-
pal; g, first phalanx; h, second
phalanx ; i, humerus ; j, ulna ; k,
accessory carpal; I, ulnar carpal;
m, intermediate carpal; n, small
metacarpal; o, bones of the ac-
cessory digit; p, third phalanx;
1, rudiment of acromion ; 2, tuber
scapulae; S, lateral tuberosity of
the humerus; 4, deltoid tubero-
sity? 5, musculospiral groove; 6,
radial tuberosity; 7, metacarpal
tuberosity; 8, tuber spinse; 9,
nutrient foramen of scapula; 10,
olecranon process ; 11, olecranon
fossa ; IS, upper interosseous
space ; IS, lower interosseous
space.
154 The Philippine Journal of Science mi
the carabao. The ulna is rather more slender and less curved
in its length.
Carpals. — The carpus consists also of six carpal bones — four
in the proximal row and two in the distal row. The bones
are very much reduced in size, otherwise they are similar to
those of the carabao.
Metacarpals. — As in the carabao, two bones are present in
the metacarpus of the timarau, the large metacarpal bone formed
by the consolidation of the third and the fourth and the lateral
small metacarpal or the fifth metacarpal bone. The large me-
tacarpal is relatively shorter, but it is not very much expanded
in its distal part, as is the case in the carabao; its tuberosity
(metacarpal tuberosity) is rather poorly developed.
Phalanges and sesamoids. — Aside from the difference in size,
the phalanges and sesamoids of the digits — the fully developed
third and fourth and the rudimentary second and fifth — cor-
respond almost exactly in general forms and characters with
those of the carabao.
THE BONES OP THE PELVIC LIMB
Os coxm. — The os coxse correspond almost exactly in general
form to those of the carabao. The following differential points,
however, are noteworthy: The crest of the ilium is almost
straight; the gluteal line is very faint; the psoas tubercle is
less pronounced ; and the tuber coxse are less tuberous and mas-
sive. The tuber ischii is likewise less massive, and the superior
ischiatic spine has fewer and less-developed vertical lines
laterally. The conjugate diameter of the anterior aperture or
inlet of the pelvis is 16.5 centimeters, while the transverse
diameter is 12.5 centimeters.
Femur. — The femur of the timarau differs only from that
of the carabao in size, being relatively shorter and less vo-
luminous, in addition to the presence of a rather deep supra-
condyloid fossa and less-developed supracondyloid crests.
Tibia. — The shaft of the tibia is less curved and the muscular
lines (linea muscular is) on the posterior surface are fewer
and less distinct. In other respects this bone resembles that
of the carabao.
Patella and fibula. — The patella and the fibula are very much
reduced in size, but in other features they correspond almost
46,1
Sumulong: Skeleton of the Timarau
155
exactly with those of the cara-
bao.
Tarsals, metatarsals, phal-
anges, and sesamoids. — The tar-
sal and metatarsal bones as well
as the phalanges and sesamoids
of the pelvic limb are likewise
almost identical with corre-
sponding bones in the carabao;
they present no striking feat-
ures except their small size.
BIBLIOGRAPHY
Bartlett, A. D. (A report on the ti-
marau.) Proc. Zool. Soc. London
(1878) 882-883.
Heude, P. M. (S. J.). Mem. Hist.
Nat. Emp. Chinois 2 (1888)
50-51.
Montellano, Pedro. The Carabao.
Philippine Education Company,
Inc., Manila (1929).
Meyer, A. B. (A report on the ti-
marau.) Proc. Zool. Soc. London
(1878) 881-882.
Miller, Merton L. The Mangyans
of Mindoro. Philip. Journ. Sci.
§ D 7 (1912) 135-156, pis. 1-10.
Peres de la Campagnie de Jesus.
Note sur le petit Buffle Sauvage
de L'ile de Mindoro (Philip-
pines). Mem. Hist. Nat. Emp.
Chinois. Imprimerie de la Mis-
sion Catholique 2 (1888) 50, 51.
Steere, J. B. (A report on the ti-
marau.) Proc. Zool. Soc. London
(1888) 415.
Sclater, P. L. The "Tamaron" of
the Philippine Islands. Nature
38 (1888) 364.
Sumulong, Manuel D. Some ob-
servations on the characteristic
features of the skeleton of the
carabao. Philip. Agr. Rev. 19
(1926) 311-325, pis. 73-81.
Fig. 4. Lateral aspect of the articulated
bones of the pelvic limb of the timarau.
a, Ilium ; b, ischium ; c, femur ; d, pa-
tella; e, tibia, /, distal end of fibula
(lateral malleolus) ; g, tibial tarsal; h,
fused second and third tarsals ; i, first
phalanx ; j, third phalanx ; k, proximal
part of fibula; I, fibular tarsal; w, fused
central and fourth tarsals ; n, large meta-
tarsal ; o, bones of the accessory digit ;
p, second phalanx ; 1, tuber sacrale ; 2,
tuber coxse ; S, nutrient foramen of the
femur ; 4, crest of tibia ; 5, greater
sciatic notch ; 6, superior ischiatic spine ;
7, tuber ischii ; 8, trochanter major ; 9,
supracondyloid fossa ; 10, nutrient fora-
men of tibia; 11, tuber calcis.
ILLUSTRATIONS
Plats 1
Fig. 1. A timarau near Bongabong River, Mindoro. (Photograph by E.
A. Heise, 1921.)
2. Timarau Bubalus mindorensis Heude, from a living animal in Me-
han Gardens, Manila. This species is restricted to Mindoro.
(Photograph by Cortes.)
Plate 2
Lateral view of the mounted skeleton of an adult timarau in the ana-
tomical museum of the College of Veterinary Science, University of
the Philippines (Photograph by the College of Agriculture.)
Plate 3
Anterolateral view of the mounted skeleton of an adult timarau of the
anatomical museum of the College of Veterinary Science^ University
of the Philippines. (Photograph by the College of Agriculture.)
TEXT FIGURES
FIG. 1. Lateral view of the skull of the timarau. a, Part of the frontal
bone; b, lacrimal; c, malar; d, nasal; e, maxilla; /, premaxilla;
g, temporal; h, occipital; i, hyoid; j, mandible. 1, horn core;
2, lacrimal fossa; 3, infraorbital foramen; U, mental foramen;
5, maxillary tuberosity; 6, temporal fossa; 7, coronoid process
of the mandible; 8, temporal crest; 9, mastoid process; 10, ex-
ternal acoustic meatus; 11, paramastoid process; 12, zygomatic
arch; IS, lacrimal bulla.
2. Frontal view of the skull of the timarau. a, Fused interparietal
and supraoccipital ; b, fused dorsal or horizontal parts of the
parietals; c, frontal bone; d, lacrimal; e, malar; /, nasal;
g, maxilla; h, premaxilla; 1, horn core (processus cornus) ;
2, external parietal crest; 3, supraorbital foramen; 4, supraor-
bital groove; 5, orbit; 6, maxillary tuberosity; 7, nasal process of
premaxilla; 8, palatine process of premaxilla; 9, palatine fissure;
10, palatine (notch) cleft.
3. Lateral aspect of the articulated bones of the thoracic limb of the
timarau. a, Scapula; b, radius; c, radial carpal; d, fused second
and third carpals; e, fourth carpal; /, large metacarpal; g, first
phalanx; h, second phalanx; i, humerus; j, ulna; k, accessory
carpal; I, ulnar carpal; m, intermediate carpal; n, small meta-
carpal; o, bones of the accessory digit; p, third phalanx; 1, rudi-
ment of acromion; 2, tuber scapulae; 3, lateral tuberosity of the
humerus; h, deltoid tuberosity; 5, musculospiral groove; 6, radial
tuberosity; 7, metacarpal tuberosity; 8, tuber spina*; 9, nutrient
foramen of scapula; 10, olecranon process; 11, olecranon fossa;
12, upper interosseous space; 13, lower interosseous space.
157
158 The Philippine Journal of Science
Fig. 4. Lateral aspect of the articulated bones of the pelvic limb of the
timarau. a, Ilium; 6, ischium; c, femur; d, patella; e, tibia;
/, distal end of fibula (lateral malleolus); g, tibial tarsal; h,
fused second and third tarsals; i, first phalanx; /, third phalanx;
k, proximal part of fibula ; I, fibular tarsal ; m, fused central and
fourth tarsals; n, large metatarsal; o, bones of the accessory
digit; p} second phalanx. I, tuber sacrale; 2, tuber coxae; St
nutrient foramen of the femur; U> crest of tibia; 5, greater
sciatic notch; 6, superior ischiatic spine; 7, tuber ischii; 8,
trochanter major; 9, supracondyloid fossa; 10, nutrient foramen
of tibia; 11, tuber calcis.
Sumulonq: Skeleton op the Timarau.]
[Philip. Journ. Sci., 46, No. 1.
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Sumulong: Skeleton op the Timarau.1
[Philip. Journ. Scl, 46, No, 1.
PLATE 3. ANTEROLATERAL VIEW OF THE MOUNTED SKELETON OF
AN ADULT TIMARAU IN THE ANATOMICAL MUSEUM OF THE COLLEGE
OF VETERINARY SCIENCE, UNIVERSITY OF THE PHILIPPINES.
The Philippine
Journal of Science
Vol. 46 OCTOBER, 1931 No. 2
RAT-BITE FEVER IN THE PHILIPPINES
By Ana Vazquez-Colet
Of the Division of Biology and Serum Laboratory
Bureau of Science, Manila
THREE PLATES
The existence of rat-bite fever (sodoku) in the Philippines
has evidently been suspected by a few practicing physicians.
Consultation of the literature reveals a clinical case reported
by Dr. Manuel Guerrero 1 and another clinical case is reported
by Jose Montes.2 As far as their reports show, their observa-
tions were purely clinical ; they did not demonstrate or identify
in the suspected cases the presence of the spirochete that causes
the disease.
The object of the present communication is to report a case
of rat-bite fever in a native child in Manila in which the causa-
tive agent of the disease, Spirochzeta morsus muris, was de-
monstrated and identified.
HISTORY OF THE ILLNESS
On January 20, 1931, about 9 a. m., a little girl, 4 years old,
named Iluminada Flores, residing at 350 Sevilla, San Nicolas,
was brought to the Bureau of Science for Pasteur treatment,
after having been bitten by a rat. The patient presented a very
conspicuous cedematous swelling of the forehead, the sides of
the face, and the neck. The sides of the face were overlaid by
distinctly visible maplike red macules. A vivid recollection of
1Rev. Filip. de Med. y Farmacia (July, 1917).
2 Rev. Filip. de Med. y Farmacia 14 (1923) 304.
262412 159
IgO The Philippine Journal of Science 1931
a graphic description of a case of rat-bite fever in Spain, re-
ported by Pascual Escolano,3 enabled me to diagnose this case
at first sight as rat-bite fever. Lt. Col. Hayashi Hirano, Med-
ical Corps, Imperial Japanese Army, at present in the Bureau
of Science, was consulted and gave me very useful suggestions
on how to proceed in isolating the causative spirochete.
The patient was bitten on the forehead by a rat on January
5, 1931, late at night, while in bed. The bite healed quite un-
eventfully within the next few days. One week after she had
been bitten, she developed fever at 6 p. m., which lasted the
whole night up to 10 a. m. the following day. She was then
free from fever for two days. Fever again appeared at 6 p. m.
and lasted three hours. This time the skin around the healed
bite, covering an area about the size of a peso coin, was swollen
and red. The patient was free from fever for two days. She
again developed fever lasting from 6 p. m. to 8 p. m. On Jan-
uary 19, 1931, at 6 p. m., the patient developed fever which
lasted two hours. In the meantime the swelling and red spots
had gradually extended to include the forehead, the sides of
the face, and the sides of the neck, but the patient evidenced
no particular discomfort, and continued to play and eat quite
as usual.
On January 20, 1931, about 9 a. m., the patient presented the
following symptoms:
1. An extensive diffuse, rather firm, oedema of the forehead
and the sides of the face and neck ; more extensive on the left
side of the neck than on the right side.
2. An elongated, irregular, maplike red macule with elevated
edges on each side of the face between the ears, posteriorly,
and the cheeks, anteriorly, and extending from the level of the
eyes above to near the edge of the lower jaw below.
3. A dusky, purplish, discolorization of the forehead was not
very conspicuous on account of the brown complexion of the
patient and the adhering remains of the ointments applied.
4. A reddish discolorization of the sides of the neck extending
from the ears and the angle of the jaw to a little above the
clavicle, on the left side, and only half-way this distance on
the right side. Purplish blotches here and there on the left side
of the neck.
5. A tiny white scar at site of bite, measuring 0.5 centimeter
vertically and 0.3 centimeter horizontally, visible on close in-
3 Rev. Med. y Cir. (1919).
46,2 Vazquez-Colet: Rat-bite Fever 161
spection, at a point situated at the junction of upper and middle
third of the forehead, somewhat to the left of the median line.
6. A marked swelling of the anterior auricular and superior
cervical lymphatic glands. A swelling of the supraclavicular
lymphatic glands on the left side was also present.
7. All the swollen parts of the face felt warm to the touch,
but the patient had no fever at that time.
8. The tongue was clean and the throat and tonsils normal.
9. The heart, the lungs, and the nervous system were normal.
January 21, 19 SI, 2. SO p. m. — The oedema and swelling of the glands
still persist. The macules are now dusky red. An injection of sodium
cacodylate combined with strychnine and sodium glycerophosphate was given
the patient.
January 22, 1981. — The patient's mother reports that the patient had
fever from 8 p. m. yesterday to 5 a. m. this morning. The tongue appears
coated and oedema persists. The edges of the erythematous areas on the
face and forehead are more elevated and are redder than the rest of
the areas. The redness on the neck is less conspicuous now, and the glands
are larger and softer. Injection of cacodylate, etc., was given.
January 28, 1981, 2.15 p. m. — The cedema has somewhat subsided. The
macules are paler red than before. The glands have reduced in size.
Cacodylate, etc., were again injected.
January 2U, 1981, 9 a. w. — The cedema has further subsided. The ma-
cules are paler and the glands smaller. A blood count was made and
the haemoglobin determined.
January 26, 1981, 2 JO p. m. — The patient is reported to have had fever
from 8 p. m. January 24 to 5 a. m. January 25 and now appears quite
pale. The macules have a light dusky reddish discolorization, and only
the edges of the macules on both sides of the face show a marked red
color. The cedema has greatly subsided; there is no redness on the neck.
The tongue is still coated. Injection of cacodylate, etc., was given.
January 27, 1981, 2 JO p. m. — The macules have practically disappeared;
only a little light red line on both sides of the face laterally to the malar
region is still distinctly visible. The tongue is clearing up and the cedema
has further subsided. The glands are smaller, but quite palpable. The
patient is pale. Cacodylate, etc., were injected.
January 28, 1981, 2 JO p. <m.— The patient is still pale, but the tongue
is now clean. The little red lines on the face are still conspicuous. The
cedema is less though the glands are still palpable. Cacodylate, etc., were
given.
January 80, 1981, 2 JO p. m.— The patient's face presents a conspicuous
appearance; a red, elevated line can be distinctly traced from a point at
the level of the lower margin of the mandible, about one inch distant
from the lobe of the ear, upward across the face to the right lower eye-
lid extending across the bridge of the nose, and following as an exactly
symmetric line on the opposite side of the face to the lower margin of
the left mandible. This line delimits symmetrical portions of the face, like
a mask, the portions above the line being a pale dusky red, quite distinct
from the portions of the face situated below the line. The cedema is
162 The Philippine Journal of Science 1931
practically gone, the glands are smaller, and the tongue is clean; caco-
dylate, etc., were injected.
January 31, 1981, 945 a. m. — The above described line on the face of
the patient is still visible, but no longer elevated. The oedema is not no-
ticeable. The glands are still palpable. Cacodylate etc., were injected.
February 2, 1931, 2 JO p. m. — The line on the face is still visible, though
not elevated, and on both sides of the face laterally to this line and about
0.75 inch distant from it, there is another red line, the intervening skin
between the two lines being pale. The mask appearance is still distinctly
visible. The glands are much smaller but still palpable.
February 3, 1931, 8 a. m. — The masklike effect is still present. The
glands are smaller.
February 4, 1931, 9 a. m. — The red line described above as delimiting
symmetrical portions of the face, has rounded up to include the lower
part of the chin. The unaffected portions of the face are now the lower
half of the nose, the inner half of the cheeks, and the tip of the chin.
This picture is quite similar to that of Escolano's case. On the affected
portions of the cheeks alternating red and pale areas of skin are seen,
the red lines describing irregular turns inclosing in some places fanciful,
flowerlike patches of pale skin. The glands, especially the upper cervical,
are still palpable. Cacodylate, etc., were given. Wassermann reaction
+ +, and Kahn reaction -f + -}-+. 4
February 5, 1931. — The masklike effect is still present, though the red-
ness is less. The red line has bridged over from under the mandible to
the left angle of the mouth. The glands are still palpable. Cacodylate,
etc., were injected.
February 6, 1931. — The red line has bridged over from under the man-
dible to the right angle of the mouth; the redness is less. The anterior
auricular glands are not palpable now, but the superior cervical glands
are still palpable. Cacodylate, etc., were injected.
February 7, 1931. — The redness has greatly faded in all the masklike
area. Cacodylate, etc., were injected.
February 9, 1931. — The masklike area is hardly visible. The redness
has disappeared; instead, a brownish discolorization now occupies the
previously reddish portions of the masklike area. The superior cervical
glands are much smaller. Cacodylate, etc., were injected.
February 10, 1931. — A few reddish lines on the cheeks and on the sides
of the chin are now seen. The superior cervical glands are still palpa-
ble. Cacodylate, etc., were injected.
February 11, 1931. — Reddish blotches are now present on the left tem-
poral region. The superior cervical glands are still palpable. Ten centi-
grams of myosalvarsan (iso) was administered intramuscularly.
February 12, 1931. — No more reddish blotches anywhere. The patient
had fever last night from 7 to 12 p. m. The superior cervical glands
are smaller.
February IS, 1931. — The patient looks well. The glands are smaller.
February 16, 1931. — The patient looks well. The glands are still pal-
pable, though greatly reduced.
*The serologic reactions were kindly performed by Dr. 0. Garcia and
read jointly by him and the author.
**>* Vazquez-Colet: Rat-bite Fever 163
JANUARY 24, 1931
Haemoglobin, per cent (Tallquist-Newcomer)5 57
Red cells per cubic millimeter 4,750,000
White cells per cubic millimeter 7,100
Differential count:
Neutrophiles, per cent 60.5
Small lymphocytes, per cent 32.0
Large lymphocytes, per cent 2.0
Mononuclears, per cent 1.5
Eosinophiles, per cent 4.0
100.0
FEBRUARY 6, 1931
Haemoglobin, per cent (Tallquist) 50
Red cells, per cubic millimeter 4,370,000
White cells, per cubic millimeter 11,500
Differential count:
Neutrophiles, per cent 68
Small lymphocytes, per cent 25
Large lymphocytes, per cent 2
Mononuclears, per cent 4
Eosinophiles, per cent 1
100
February 6, 1931. — Reaction acid; glucose negative, albumin traces. Se-
diments: Abundant epithelial cells, leukocytes and amorphous urates, few
mucous threads and cylindroids. No casts found.6
PROCEDURE EMPLOYED TO DEMONSTRATE THE CAUSATIVE AGENT OF
RAT-BITE FEVER
A few drops of blood were obtained from the patient's fore-
head at points near the site of the bite and inoculated intraperi-
toneal^ into a white mouse (RB-Ms-1), and subcutaneously
into the abdomen of a guinea pig (RB-M-1) . Some gland juice
was obtained from one of the superior cervical lymphatic glands
(left side) and injected subcutaneously into the abdomen of a
white mouse (RB-Ms-2).
Smears from tissue scrapings, obtained by scraping two inci-
sions made near the site of the bite, were also prepared January
20, 1931. The smears were stained by Giemsa's method. The
spirochete was demonstrated in them.
Erythrocyte and leukocyte counts and haemoglobin percentage deter-
minations were kindly made by Dr. Jose* Ramirez.
6 Routine examination of the patient's urine was performed by Dr. G.
Sepulveda, Jr.
164
The Philippine Journal of Science
1931
The blood of the mice and guinea pig was examined daily by
dark-field illumination. Mouse RB-Ms-2 showed spirochetes
in its blood for the first time after inoculation January 30,
1931; that is, ten days after its inoculation. Mouse RB-Ms-1
became positive February 5, 1931; that is, sixteen days after
it was inoculated. The spirochetes were demonstrated in the
blood of the mice both by dark-field examination and in smears
stained by Giemsa's method. February 5, 1931, some peritoneal
fluid from the mice was obtained, using fine capillary tubes.
Smears were prepared and stained by Giemsa's method. A few
spirochetes were demonstrated.
February 7, 1931, the guinea pig showed palpable inguinal
lymphatic glands. February 9, 1931, the glands were larger,
especially on the left side. From one of these some gland juice
was obtained by means of a fine capillary tube and examined
both by dark field and in smears stained by Giemsa's method.
No spirochetes were seen by dark-field examination, but in the
stained smears several spirochetes were seen. This was twenty
days after the guinea pig was inoculated.
February 10, 1931, twenty-one days after inoculation, some
peritoneal fluid was obtained from the guinea pig and exam-
ined both by dark field and in smears stained by Giemsa's method.
Microns.
Coils.
Spirochetes from blood of RB-Ms-2 *
2.0
2.5
2.5
2.5
2.3
3.6
3.8
4.98
3.7
2.5
3.0
2.5
3.3
2.4
3.3
3.3
2.0
3.3
2.5
2.5
2.5
3
4
5
5
6
7
7
9
7
4
5
4
6
4
5
5
3
7
3
4
5
Do
Do _ _
Do _
Do
Spirochetes from blood of RB-Ms-1
Do _
Do _ _
Do _
Spirochetes from tissue scrapings of patient
Do _
Do
Spirochetes from gland juice of guinea pig RB-M-1
Do -
Do
Do __ _. __ .
Do __. _
Spirochetes from blood of guinea pig RB-M-1
Dc__ _
Do _
Do _
8 Measurements of the spirochetes were kindly taken by Dr. Marcos Tubangui jointly
with the author.
46,2 Vazquez-Colet: Rat-bite Fever 165
No spirochetes were detected by dark field, but in the stained
smears spirochetes were seen. It was not until February 15,
1931 — that is, twenty-six days after inoculation — that the guinea
pig showed the spirochetes in its blood. They were demon-
strated in smears stained by Giemsa's method.
MORPHOLOGY AND MOTILITY OF THE DEMONSTRATED SPIROCHETE
Under dark-field illumination the spirochete was seen as a
rather short and rigid spindle-shaped organism, which darted
back and forth very quickly and as quickly disappeared by
shooting to one side. The spirochetes could be seen in the
clear spaces between the red blood cells. The organisms moved
so fast that details of their structure could not be observed.
Only occasionally, when the spirochete came to rest, could it be
seen that its body is undulated, the undulations apparently lying
in one plane. In stained smears the organism was seen to be
much the same as Vandyke Carter describes it (called by him
Spirillum minus) ; namely, that it is "an extended and uniformly
slender filament of clearly spiral construction, having a length
commonly somewhat less than the diameter of a blood disc but
varying from 5 microns to 9 microns, and according to its length
presenting from four to eight close spiral turns."
CONCLUSIONS
1. A clinically typical case of rat-bite fever was accidentally
encountered among cases reporting for antirabic treatment.
2. The causative agent of rat-bite fever, Spirochzeta morsus
muris, was demonstrated in tissue smears from the patient and
by inoculating it into experimental animals and recovering it
from them by microscopical slides, stained and dark field. The
spirochete was identified morphologically and by measurements
as well as motility to be Spirochseta morsus muris.
3. Thus the existence in the Philippine Islands of rat-bite
fever was definitely established.
ACKNOWLEDGMENTS
To Dr. Otto Schobl, chief of the division of biology, I wish to
express my appreciation for suggestions offered.
To Lt. Col. Hayashi Hirano, Medical Corps, Imperial Japanese
Army, now detailed at the Bureau of Science, I am also indebted
for his cooperation in demonstrating the presence of the parasite.
ILLUSTRATIONS
Plate 1
Fig. 1. Front view of the patient's face showing the site of the bite on
the forehead and the lesion that developed at the site of the
bite.
2. Showing the swollen cervical glands.
Plate 2
Side view of the patient showing oedematous feature of the lesion on the
forehead and the swollen cervical glands.
Plate 3
Smears stained by Giemsa's method. Photomicrograms taken with ocular
No. 4 and 1/12 oil immersion.
Figs. 1, 2, and 4. Showing different sizes of Spiroch&ta morsus muris and
its relation to the size of the red corpuscles.
Fig. 3. Two spirochetes joined end to end.
167
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PLATE 2.
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PLATE 3.
AN INTERPRETATION OF THE LAWS OF BROWN AND
PEARCE THAT GOVERN THE COURSE
OF TREPONEMATOSES *
By Otto Schobl
Chief of the Division of Biology and Serum Laboratory
Bureau of Science, Manila
From the results of their classic experiments with syphilis on
rabbits, Brown and Pearce deduced two laws that regulate the
biologic events taking place during experimental infection and
that are directly traceable in the course of natural infection in
man. At least one of these laws was found by Schobl1 to
apply to yaws as well, with a slight modification, conforming to
the biologic nature of the parasite of yaws as it differs from
that of the parasite of syphilis. Clinical observation showed
that the law, which holds true in experimental yaws in animals,
likewise applies to natural or experimental yaws infection in
man. The two are known as the law of inverse proportions and
the law of sequence.
The law of inverse proportions, as it applies to both syphilis
and yaws, says: The more intensive the early manifestations,
the less intensive are the late manifestations of the disease.
This law, as expressed above, applies as much to syphilis as it
does to yaws. In syphilis, however, it enters into play in the
relation between the primary and the secondary stage, as well
as between the secondary and the late manifestations; while in
yaws it is true only between the early stage on the one hand,
and the late manifestations on the other.
The second law of Brown and Pearce is the law of sequence.
The various systems of tissues are affected successively. This
law is clearly evident in human syphilis where the integument,
the internal organs, the cardiovascular system, and the central
nervous system are affected successively and in combinations
with great regularity. In yaws this law has little application
due to the epidermotropic tissue selectivity of the parasite that
* Received for publication February 5, 1931.
1 Philip. Journ. Sci. 35 (1928) 211.
169
170 The Philippine Journal of Science 1931
causes this disease. While early yaws lesions are restricted to
the skin exclusively, the late ulcerative lesions also occur in the
skin but may by continuity migrate into the tissues immediately
attached to the integument. Thus subsequent to an ulcerative
skin lesion the muscle, the periost or cartilage, and even the bone
may be affected by an hypertrophic, atrophic, or ulcerative pro-
cess originating in the skin. By the time the lesion is seen in
the clinic the original skin lesion may have healed by scars while
the lesion in the bone, for instance, may persist at the time when
the patient is first seen. It is nothing but a part of the original
skin lesion that migrated, healing as it traveled. The entire
course of development of such lesions is never seen in the clinic.
The clinician has entered the theater in the third or last act of
the drama. Unless these lesions are experimentally produced
and followed step by step, the pathogenesis of late yaws lesions
remains an unsolved mystery to the clinician, who is surrounded
on a yaws clinic by a veritable kaleidoscope of chronic clinical
phenomena, the past and the future of which may never come
within the range of his vision.
The present author is unaware of an adequate interpretation,
or any at all, of the laws as first formulated by Brown and
Pearce. In the course of experimental work on yaws and sy-
philis, performed partly on human volunteers and partly on
suitable animals, which the present author has carried on in
the course of the last seven years, certain findings were made
that correlated themselves, as the work progressed, into a logical
chain of what appeared to be natural causes of the nosologic
phenomena that form the clinical course of treponematoses and
for which laws were deduced by Brown and Pearce.
The first observation in this direction was made when it was
found that the intensity of early yaws lesions stands in direct
proportion to the number of treponemas contained therein.2
Since the intensity of the early lesions stands in inverse pro-
portion to the intensity of the late lesions, in yaws as well as in
syphilis, the law could be expressed thus: The relative number
of invading parasites in the early stage of infection stands in
inverse proportion to the intensity of the late manifestations.
The second finding was that the serologic response due to
superinfection stands in inverse proportion to the serologic
response of the original infection.3 The law of inverse propor-
2 Philip. Journ. Sci. 35 (1928) 257.
8 Op. cit. 272.
46> 2 Sehobl: Laws of Brown and Pearce \q\
tions was found reflected in the serologic picture of experimental
yaws and syphilis.
The next link in the chain of experimental results was the
finding that a time relation exists between the late serologic
response and immunity.4 At the time the late response be-
comes apparent in the form of strong serologic reactions the
resistance to superinoculation is fully developed, and any ex-
perimental procedure that accelerates the late serologic re-
sponse hastens the development of resistance to superinoculation
or reinfection. From our early experiments with yaws it is
known that no new lesions form, either secondary or late ulcera-
tive ones, from the time the yaws monkeys become immune to
superinoculation; but the respective generalized or late ulcera-
tive lesions that have developed before that time persist. We
formulated this finding in the statement: The time during which
the secondary or the late ulcerative yaws lesions form is limited
by the development of immunity and is shorter than the time
necessary for the healing of the already existing lesions. Prior
to the onset of such a high degree of immunity that it completely
prevents the formation of specific lesions at the place of super-
infection, metastatic lesions develop that are atypical and have
been called by us framboesides. Late ulcerative lesions may
form at the place of superinfection at this time.5 Both types
of lesions contain such a small number of treponemas that it is
difficult and frequently impossible to demonstrate their presence
in the lesions by dark-field microscope. These lesions occur
after the typical ones and before complete resistance sets in.
From these experimental findings we have deduced the explana-
tion that partial immunity is responsible for the modification of
the morphology of treponematous lesions.
The law of inverse proportions, which is the first law of
Brown and Pearce, can be expressed as follows : The number of
the invading treponemas during the early stage stands in direct
proportion to the degree of immunity that subsequently develops.
It stands in inverse proportion to the time necessary for the
development of immunity, or in other words, the number of
invading parasites in the early stage of infection stands in
direct proportion to the speed with which immunity derelops,
the speed being the ratio between quantity and time. This law
of direct proportions between the number of invading parasites
4 Philip. Joum. Sci. 42 (1930") 203; 43 (1930) 603.
5 Philip. Journ. Sci. 35 (1928) 230-236; 242-251.
172 The Philippine Journal of Science 1931
and the speed of development of consequent immunity applies
not only to the living parasites but also to the lifeless antigen,
to the infection and the following vaccination or vice versa.
Thus the law has a general application and can be finally formu-
lated as follows: The degree of subsequent immunity and the
speed of the development of immunity stands in direct propor-
tion to the amount of treponematous antigen. Due to this di-
rect proportion between the treponematous antigen and the sub-
sequent immunity, the immunity stands in indirect proportion
to the duration of the clinically active disease.
The law of sequence indicates successive involvement of va-
rious tissue-systems by the syphilitic infection. The trepone-
mas invade the blood stream from the initial portal of infec-
tion in its early stage. This is true of syphilis as well as of
yaws. In the case of yaws, contrary to syphilis, the parasites do
not colonize the internal organs permanently and do not produce
lesions in these organs. In the case of syphilis, the parasites
remain viable in the mesoderm for a very long time, if not for
life.
The treponemas being disseminated into the various tissues
through the blood stream in the very early stage from the portal
of infection, the law of sequence is not based on a successive
invasion of the various tissues by the parasites. The integu-
ment comes in contact with the treponemas first of all. They
invade the internal organs in the early stage of infection, but
the heaviest immigration into these organs takes place when
the treponemas are present in the largest numbers in the initial
lesion; that is, at the time when the initial lesion is fully de-
veloped. Thus the tissues of the integument, which form the
seat of the initial lesion, in a typical clinical case of syphilis
pass from the stage of sensitization through the negative phase
into the positive phase somewhat ahead of the other tissues.
The immunity is transferred from one to the other systems of
tissues; first in the form of a delayed incubation period, then
in the form of a changed clinical and anatomical morphology of
the lesions. It is very likely true of all infections, but in
treponematoses, and particularly in syphilis, it is clinically
evident that before immunity becomes fast a more or less
pronounced oscillation between the negative and positive phase
of immunity takes place. This swinging of the pendulum be-
tween the positive and the negative phase is not necessarily
synchronic in all tissues, because even in normal skin the in-
cubation period of two or more experimental lesions produced
46ȣ Schobl: Laws of Brown and Pearce 173
by simultaneous inoculation of the same amount of the same
yaws-inoculum, by the same method, and under the same tissue
conditions, even in symmetric parts of the same animal, is not
always the same. Thus the fate of a focus of treponemas de-
posited in a given tissue, in the course of treponematous in-
fection, is influenced by the phase of the transmitted immunity
due to the activity of another focus of treponemas deposited
in another place of the same or in another system of tissues.
An almost healed yaws lesion was brought to an extensive ex-
acerbation, and dormant deposits of yaws treponemas were
incited to formation of lesions, after an extraordinarily long
incubation, by superinoculation with syphilis that failed to
produce syphilitic lesion. On the other hand superinoculation
of yaws monkeys with syphilis that produced syphilitic lesion
resulted not in exacerbation of the existing yaws lesion but in a
striking acceleration of yaws-immunity.6 This shows that im-
munity in its negative as well as in its positive phase is trans-
mitted between tissues.
An immunity that is on an upgrade incline may be accelerated
into the positive phase, while the immunity in another part of
the same system of tissues or in different systems of tissues that
is on the downward incline may be accelerated into a deeper
negative phase by superinfection than would be otherwise pos-
sible. It is clearly evident that spontaneous exacerbation of a
lesion takes the place of an experimental superinfection in this
respect. One is a superinfection from within, the other from
without. The decisive factor is the sudden increase of trepone-
matous antigen, dead or alive, that is brought in contact with
mesodermic tissues in these instances. The variation of the in-
cubation period, which is made much more elastic by the initial
immunity, makes incalculable the possible effects of the inter-
mingling immunity-phases on the course of the main immunity
curve, which in itself is not steady. They can, however, be
predicted in a general way.
The second law in syphilis of Brown and Pearce, that is, the
law of sequence, is here interpreted as a sequence of immunity
that develops successively in the various systems of tissues.
The treponemas that invade the various tissues, long before the
immunity has developed, can multiply and produce lesions only
in those tissues that are not yet immune. Not all tissues are
equally capable of producing immunity in treponematoses. This
•Philip. Journ. Sci. 42 (1930) 239.
174 The Philippine Journal of Science mi
is evident from the findings that treponema pertenue introduced
into the epiderm causes immunity to develop in six months;
when introduced into the mesoderm in six to eight weeks.T
The explanation given here, that is, the successive transmis-
sion of immunity from tissue to tissue, explains the well-known
clinical observation that specific syphilitic lesions develop in the
internal organs or in the central nervous system in a host whose
integument has long become immune to reinfection and to
relapses. A better explanation of the condition known as neu-
rosyphilis is to assume that due to insufficient immunization in
the early stage of the infection, which may have been caused by
mild early lesions, by insufficient sterilization of the host, by
treatment given in the early stage of the disease, or by super-
infection, taking place in a partially immune body, it may have
assumed the symptomless form, rather than the explanation
given at times in the literature that, due to the modern arsenical
treatment, the syphilitic infection becomes neurotrophic. Strains
of treponema pallidum isolated from neurosyphilis produce ty-
pical chancres in experimental animals and otherwise behave
like any other strain isolated from a primary lesion. They do
not show any signs of permanent changes in their biology and
behave differently in the body of the neurosyphilitic case, from
which they were isolated, on account of the changed condition
of that particular patient's tissues and not on account of a
change in the biology of the parasites. A simple experiment
convinced us of the truth of our supposition that immunity in
syphilis involves the various tissues at different stages and in
succession. A series of yaws monkeys inoculated with yaws
more than a year prior to this experiment, and that repeatedly
had been proven immune to yaws were inoculated with Nichols
strain of syphilis on one side of the scrotum intradermally .
No lesion developed at the place of inoculation but the normal
control animal developed a typical sclerosis. In due time the
corresponding inguinal glands were excised and transferred to
rabbits' testicles. None of the rabbits that received the glands
from the immune monkeys developed lesions and they were
found susceptible to syphilis five months later. Thus it was
proven that the lymph glands contained no treponemas. The
rabbits that received the lymph glands from the nonimmune
controls developed typical chancres. Thus it was proven that
the inoculum contained viable virus of syphilis. Two months
7 Philip. Journ. Sci. 35 (1928) 280; 45 (1931) 221.
46,2 Schobl: Laws of Brown and Pearce 175
later the immune monkeys were reinoculated with syphilis by
intratesticular injection on the opposite side from the point
where the first inoculation with syphilis was introduced. The
lymph glands, corresponding to the place of the second, the in-
tratesticular inoculation, were transplanted to rabbits. One
half of the rabbits developed chancres and the other half re-
mained normal. The latter animals were found susceptible to
syphilis five months later. This experiment shows that all of
the animals immune to yaws were also immune to syphilis as
far as skin was concerned, but only some were immune to sy-
philis with regard to internal organs at that time. Two months
after the skin immunity was established the internal organs
concerned were immune only in some of the experimental animals
and not in others.
SUMMARY
The law of inverse proportions of Brown and Pearce is inter-
preted as a direct proportion between the quantity of trepone-
matous antigen, dead or alive, and the degree of immunity and
the speed of its development. The law of sequence of Brown and
Pearce is interpreted as a successive development of immunity
in the various systems of the body's tissues.
This interpretation, based on experimental facts, brings these
laws in agreement with the laws that govern antibacterial im-
munity, and is a contribution to the knowledge of tissue im-
munity.
262412 2
COEXISTENT INFECTION WITH YAWS AND SYPHILIS
By Otto Schobl
Chief of the Division of Biology and Serum Laboratory
Bureau of Science, Manila
Clinical observations made on man and experimental expe-
rience with humans and animals, show that yaws infection can
supersede that with syphilis and vice versa. This phenomenon
has been interpreted in various ways in the past. Those authors
who maintained that yaws and syphilis are distinct and differ-
ent diseases used these observations as proof of their dualistic
interpretation of the treponematoses, while the opposite side
claimed that lesions were found in yaws patients that could not
be distinguished clinically from syphilitic lesions and deduced
that yaws and syphilis are one disease.
The new designations that are given by the unitarians to
yaws such as "tropical syphilis/' "primitive syphilis," and
"rural syphilis," to distinguish yaws from "civilized syphilis,"
or "city syphilis," prove ipso facto that all is not well with
the unitarian theory or else such differentiation in designations
would not be necessary. It appears from the literature that the
possibility of coexistence of the two diseases in one and the
same patient, and particularly the influence that coexistent infec-
tion with syphilis may have on the clinical course and manifesta-
tions of yaws or vice versa, have not been considered.
Unfortunately, the question of the relation between yaws and
syphilis is a far deeper problem than merely one of clinical
nomenclature. It is a question of different organotropism of
the treponema of yaws from that of the treponema of syphilis.
It is not an isolated phenomenon and finds its analogy in certain
relations of leprosy to tuberculosis, of herpes to encephalitis,
and probably of dengue to yellow fever. The question of orga-
notropism with regard to infection and immunity very likely
has more general application than is suspected today, and the
once inviolable laws of specificity of infection and immunity are
being modified constantly, as is also our conception of immunity.
177
178 The Philippine Journal of Science idsi
The experimental evidence that has come to light through
our researches,1 which shows that reciprocal immunity exists
between yaws and syphilis, does not prove that the two diseases
are one and the same, as it appears to at first sight. On the
contrary, the difference in immunologic conditions existing in
yaws and in syphilis, both in animals and humans, as well as the
difference in the behavior of the two infections with regard to
cross immunity, shows plainly that fundamental immunologic
differences exist between yaws and syphilis. These differences,
like those of the tissue selectivity of the respective parasites, the
pathology, pathogenesis, clinical course, transmission, geogra-
phic and age distribution, stand in complete agreement with the
fundamental biologic distinction between the parasite that causes
yaws and the parasite that causes syphilis.
Experimental evidence shows that infection with syphilis may
have a decided effect on the course of a coexisting yaws infection.
This effect is evident in two directions. According to the stage
of immunity that is present at the time when the syphilitic lesion
develops in a yaws-infected host the immunity may swing into
a negative phase and exacerbations or relapses of the basic in-
fection may occur.2 On the other hand, the immunity may
swing rapidly into the positive phase3 and the effect of such
cross superinfection will be beneficial to the host, inasmuch as
the rapidly accelerated immunity prevents the development of
further stages of both yaws and syphilis.
Therefore, it is quite evident that syphilitic lesions, as well
as yaws lesions, may coexist in one and the same host just as
leprous lesions and tuberculous lesions may coexist in the same
patient. This coexistence certainly does not justify the con-
clusion that syphilis and yaws are one and the same disease. If
lesions that cannot be differentiated anatomically from syphi-
litic lesions are found in internal organs of yaws patients, such
as the cardiovascular system or the placenta, there is every
reason to assume that these lesions are of syphilitic rather than
of framboesic origin, and the possibility of a double infection
must be considered in such cases. There is hardly a corner of
the world where syphilis has not been introduced. A statement
made in German literature that Nichols strain of yaws, after
repeated passages through rabbits over a period of two years,
Philip. Joum. Sci. 42 (1930) 203, 239; 43 (1930) 263, 429, 583; 45 (1931)
221.
a Philip. Journ. Sci. 42 (1930) 245.
3 Op. cit. 241.
46,2 Schobl: Coexistent Infection 179
changed its character suddenly so as to become indistinguish-
able from that of syphilis, merely shows, provided that no error
was committed since both Nichols strain of yaws and Nichols
strain of syphilis have been imported to Germany, that the ex-
perimental animal used, the rabbit, is unsuitable for the study
of yaws. The subject to be studied, yaws, became unrecog-
nizable in this kind of animal. Retroinoculation to men or to
a Philippine monkey would be the only procedure in such a case.
The frequent and unduly exaggerated statements that yaws
lesions cannot be differentiated from syphilitic lesions clearly
prove that mere clinical inspection, unsupported by other meth-
ods and procedures of biologic investigation, has its limita-
tions, which must naturally vary with the dermatologic train-
ing and experience of the observer.
The crucial test that decides whether a given doubtful lesion
is of framboesic etiology is the inoculation of the material ob-
tained from this lesion to a suitable experimental animal. Thus,
an atypical yaws lesion in a patient, with which the diagnostician
may not be acquainted, is reduced to a typical initial lesion of
yaws in a suitable animal and may be easily recognized even
by a less trained or less experienced physician. Due to the great
morphologic similarity of Treponema pertenue and Treponema
pallidum the mere microscopic demonstration of treponemas in
smears or sections cannot settle whether a given lesion is of
framboesic or syphilitic origin. By inoculation of the material
from atypical yaws lesions to monkeys, we were able, on several
occasions, to confirm our clinical diagnosis of yaws and to con-
vince the attending physician that the lesion was yaws and
not syphilis.
It must be borne in mind that the treponematoses are chronic
infections, that the immunity develops slowly, that there is a
great number of possibilities in the scale of immunity from
complete susceptibility to complete immunity, and that the
quantity of early infection affects the progress of the immunity
in direct proportion as to degree and in inverse proportion as
to time.
The possibilities are further augmented by the mutual inter-
ference of cross immunity between yaws and syphilis. The
degree of immunity at a given time in the course of a trepone-
matous infection has a deciding effect on the modification of
subsequent clinical lesions. Not only the homologous but also
the cross immunity between yaws and syphilis modifies mutually
the clinical character of the lesions and the course of the dis-
180 The Philippine Journal of Science 1931
eases. This modification varies according to the degree of im-
munity existing at a given time. A treponematous lesion de-
velops when the parasites propagate at a given site in the host's
body tissues. Immunity of low grade restricts the propagation
of treponemas lodged in the tissues and a modified or atypical
lesion may develop. The highest grade of immunity suppresses
completely the propagation of the parasites in the tissues and
no lesion develops. This is true of homologous as well as of
cross immunity between yaws and syphilis. When a host in-
fected with yaws develops no lesion at all at the place of homol-
ogous superinoculation, the infection is brought to a stand-
still and no new yaws lesions will develop. The host has
reached a high degree of homologous immunity. At that time,
however, he has not yet become immune to cross infection, and
if originally infected with syphilis, for instance, may contract
yaws, with either a typical or modified course, for some time
after the superinfection with syphilis no longer produces a
lesion. From this it follows that simultaneous or subsequent
cross infection with yaws and syphilis is to be considered as a
probability in a clinical case. A cross infection is possible be-
yond the time when a high degree of homologous immunity has
developed and up to the time when a group immunity develops,
which then includes not only homologous but also heterologous
treponematous infection as well. The condition of tissue non-
reactivity that develops in syphilis after the infection has lasted
for some time is frequently referred to as anergy. This term,
however, is misleading as to the nature of this tissue condition,
insinuating, as it does, a property of tissues that has been lost
rather than a property that has been gained. Animals in-
fected with syphilis in our experiments soon reached a stage
in which a subsequent homologous superinfection no longer
produced a lesion. The absence of lesion at the place of su-
perinfection is due to the inability of the parasites to multiply
and the non-development of the lesion is due to this factor and
not to the inability of the tissues to react. The parasites do
not multiply under these conditions and consequently do not exert
sufficient irritation to cause the tissues to react. When the very
same syphilitic animals that failed to develop lesion as a conse-
quence of superinfection with syphilis were superinfected with
yaws, the tissues reacted promptly and a yaws lesion developed,
showing that the tissues were capable of reacting to the in-
troduction of parasites for a considerable time after they no
longer reacted to the homologous superinfection and before
46,2 Schohl: Laws of Brown and Pearce 181
the cross immunity to yaws developed in syphilitic animals.
There seems to be no reason why this phenomenon of so-called
anergy should not be interpreted as immunity, which suppresses
the propagation of the parasites either completely or at least
partially.
The clinical result of cross infection with yaws and syphilis
may be either an exacerbation of the basic, as well as of the
subsequent infection, or the subsequent cross infection may
bring about a partial or complete suppression of both coexisting
syphilis and yaws, in which case further stages of both diseases
will be limited or completely suppressed. Which of the two
possibilities will happen depends on the degree of immunity pres-
ent at a given stage of the disease.
THE PROSPECTS OF VACCINATION AND VACCINE
THERAPY IN TREPONEMATOSES
. *
By Otto Schobl
Chief of the Division of Biology and Serum Laboratory
Bureau of Science, Manila
The clinical course and the ultimate result of treponematous
infections, like those of any other infection, are determined by
the biology of the parasite and by the immunity that develops
in the course of the infection. Whether the innate nature of a
parasite is to multiply consecutively or intermittently, that is
in a cycle, the parasites should propagate progressively in the
body of the host and the inevitable consequence thereof should
be the death of the host. This, however, is not always the case
even in the most acute infections and it rarely occurs in such
chronic ones as the Treponematoses. The chain of subsequent
biologic events that take place in the course of treponematous
infections, which are the result of mutual interaction between
the parasite and the tissue response of the host, may be spoken
of as immunity in "statu nascendi." The ultimate immunity
prevents the parasites that have invaded the tissue of the host,
previous to or subsequent to the full development of immunity,
from further propagation. No new lesions develop from that
time on. Definite and well-known laws govern the clinical
course of treponematoses from the beginning of the infection to
the end. These laws are determined by a quantitative relation
between the parasites and immunity. There exists a* direct
quantitative proportion between the number of parasites pres-
ent in the body of the host and the degree of subsequent im-
munity, and an inverse proportion between the number of para-
sites and the time necessary for the development of immunity.
The more parasites there are present in the early stage of the
infection, the higher the degree of immunity that will develop,
and the quicker it will set in. The clinical course of the infec-
tion in treponematoses and its consequences are determined in
* Received for publication February 5, 1931.
183
184 The Philippine Journal of Science i^u
the early stage of the disease by the relative number of parasites.
The earlier the immunity develops, the shorter and less tragic
is the course of the disease. Further progress of the disease
is halted in any stage of the disease whenever full immunity
sets in. The immunity in treponematoses can be accelerated by
artificial means, and the course of the disease can be influenced
thereby. A high degree of immunity can be made to set in
before the expiration of the incubation period of the primary,
or of the generalized so-called secondary manifestations or of
the late forms.
The experimental evidence on which these statements are
based has been published, likewise the possibility of preventive
vaccination and vaccine therapy in treponematoses has been de-
monstrated on animals. It is the object of this communication
to discuss the mechanism of these vaccination procedures in
order that an appraisal of the practical possibilities may be
realized.
The intimate relation between the serologic response of the
infected body organism and the stages of immunity in statu
nascendi in the course of treponematous infections is unques-
tionable. Coincidently with the development in animals of the
primary lesion, which is the clinically visible sign of sensitiza-
tion, and provided that only local lesion develops in the course
of the infection, the curve that registers the results of serologic
examinations rises to a more or less high point, the strength
of the serum-reactions being directly proportional to the in-
tensity of the lesion, in other words to the number of parasites
present in the primary lesion. With the healing of the primary
lesion, the serologic curve returns to normal, only to rise again
to the highest point at the time when a high degree of immunity
sets in.
In animals immunized with lifeless treponematous antigen,
only the early serologic response becomes evident, that is, the
one which is coincident with the primary lesion in case of in-
fection. The late response is absent in case of vaccination with
killed treponemas, or it is possible that it is very much delayed.
If generalized manifestations appear, following the develop-
ment of the initial local lesion, the late serologic response as well
as the development of immunity is accelerated. An analogous
phenomenon occurs if infection takes place following vaccina-
tion with killed treponemas, that is, a sudden rise of the serologic
46,2 Schobl: Vaccination in Treponematoses 185
curves and acceleration of the development of immunity. We
may express it in the following way. From the serologic and
immunologic standpoint, the preventive vaccination takes the
place of the primary lesion, and the infection that follows the
preventive vaccination takes the place of the generalization of
the treponematous infection, the so-called secondaries. Clin-
ically speaking, there are several possibilities when treponema-
tous infection invades the vaccinated body organism. The result
depends on the time relation between the incubation time of the
infection and the speed of the acceleration of immunity as a
consequence of the infection itself. If the time necessary for
the accelerated immunity to reach a high degree is shorter than
the incubation of the infection in the vaccinated body organism,
then no lesion develops. If the time required by the immunity
to be accelerated to a high degree by the subsequent infection
is longer than the time of incubation, a local primary lesion
develops, but the immunity is raised thereby, to a high grade,
before the time when the generalized manifestations (the sec-
ondaries) or the late forms (the tertiaries) can occur. Con-
sequently, following the primary lesion, no subsequent stages
of the disease develop.
Specific antitreponematous treatment, when administered in
the early stage of a primary lesion, delays the onset of im-
munity far beyond the time at which immunity sets in if infec-
tion is allowed to run its course without treatment. Under
these circumstances treponematous reinfection or relapses, if
the treatment was not complete, are possible for a long time.
Vaccine therapy, administered after early specific cure, accel-
erates the immunity. Within a short time after the vaccine
therapy has been administered a reinfection is no longer pos-
sible. The primary lesion is accompanied by a rise in the serol-
ogic curve, which drops to zero following the treatment. The
development of immunity is delayed by early treatment, and the
earlier the treatment is administered the more is the immunity
delayed. In other words, a primary lesion whose progress has
been terminated by early specific treatment has the same serol-
ogic and immunologic effect that vaccination with killed trepone-
mas has in healthy animals. Intramesodermal incorporation of
antigen, living or dead, causes a rapid development of immunity.
Thus reinfection after early cure may be prevented by vaccine
186 The Philippine Journal of Science 1931
therapy, or in case that primary lesion develops due to rein-
fection no further stages of the disease -will develop.
Vaccine therapy administered to an infected host without pre-
vious or simultaneous specific antitreponematous treatment pro-
vokes a negative phase and severe lesions may appear in such a
case.
The most opportune time for an effective application of anti-
treponematous vaccination, either before natural infection has
taken place or after the body organism has been infected, is the
stage of normal or the stage of exaggerated tissue reactivity.
When the tissues reach the stage of diminished reactivity the
effect of the antigen supplied by vaccination will be slight or nil.
The immunity in treponematoses runs the general course of a
saturation curve. It follows inevitably from the shape of such
a curve that its rise can be influenced effectively in the initial
phase, somewhat in the middle phase, and very little, if at all,
in the last phase of the curve. In order to be effective the vac-
cination with killed treponemas must take the place of a vigorous
infection of the early stage so that, according to the law of
inverse proportions of Brown and Pearce, the subsequent stages
of the disease are very mild or do not manifest themselves at
all. Whether administered to a healthy body or to a previously
infected host the antitreponematous vaccination is a preventive
measure and tends to hasten an immunity that, in turn, prevents
the development of subsequent stages of the disease. It has no
apparent healing effect on the lesions that have already devel-
oped. The vaccination with killed treponemas is a controllable
and harmless substitute of a severe early treponematous infec-
tion that, as experimental evidence and clinical observation
show, prevents the development of late stages of the disease.
These statements with regard to vaccination and vaccine ther-
apy are based on experimental evidence and refer to experi-
mental animals.
A fair estimate can be made of the possibilities of vaccination
and vaccine therapy in treponematous infections in man. This
estimate has not been made by merely applying the experimental
findings made in monkeys to man, but by drawing a comparison
between the conditions experimentally found in monkeys and
the conditions that were found in experimentally inoculated hu-
mans. No rational objection can be held against such compa-
rison, particularly in the case of yaws, since it has been proven
that by appropriate experimental procedure an infection can be
produced in monkeys that runs the same course and manifests
46»2 Schobl: Vaccination in Treponematoses 187
itself in these animals in the same principal forms as does the
disease in human beings.
The incubation period of the primary yaws lesion, and of the
generalized so-called secondary stage in monkeys, is the same
as was experimentally found in humans. In monkeys the im-
munity sets in earlier than it was found to take place in ex-
perimentally infected human volunteers. This finding explains
the fact that the period during which generalized and late yaws
lesions crop out in monkeys is shorter than in man. From this
comparison, it is safe to predict that the effect of vaccination
in human treponematoses will not be as prompt as in experi-
mental animals. The degree of immunity, however, in these
infections, as well as the rapidity of its development, depends
on the amount of vaccination and on the proper time at which
immunization is carried out. Furthermore, experimental in-
oculation is a far more severe test for immunity than the one
that is usual in the great majority of natural infections of man.
We may, therefore, rightfully hope that this procedure will be
a valuable addition to our present armament for combating
human treponematoses, that is yaws and syphilis.
DECAY OF WOOD IN AUTOMOBILES IN THE TROPICS
By C. J. Humphrey
Mycologist, Bureau of Science, Manila
TWO PLATES
The depreciation of automobiles in the Tropics from decay of
the wood used in their construction reaches a staggering figure
in proportion to the investment. The various kinds of timber
used in their manufacture are not, as a rule, adapted to with-
stand the warm humid weather to which the cars are constantly
subjected. As a result of this the timber replacement business,
in the City of Manila alone, has become a considerable industry.
Most of the cars in the Philippines are of American manu-
facture ; a few come from Europe. The woods used are almost
exclusively native temperate-zone species selected for their
strength, toughness, workability, etc., rather than for their
durability. In American cars x ash, beech, birch, elm, hickory,
maple, oak, and sycamore are commonly employed in places re-
quiring strength. Hickory is regularly used for spokes and
rims; oak for top bows. For running boards, seat risers, seat
lids and other parts not requiring strength, chestnut, gum, tu-
pelo, yellow pines, Douglas fir, etc., are commonly employed.
With the exception of white oak, chestnut, resinous yellow pine,
and heart red gum these are all recognized as being only mode-
rately resistant to decay. When sapwood is used it is very
perishable regardless of the durability of the heart.
The automobile and truck business in the Tropics has reached
a considerable volume, the registration 2 for 1929 in the Philip-
pine Islands being 21,341 passenger cars and 10,365 trucks, of
which 9,545 cars and 2,965 trucks were registered for the City
of Manila. The turn-over of used cars is comparatively large,
this being to an appreciable extent occasioned by the rapid dete-
rioration of the wooden parts. Many people prefer to dispose
of a car that has seen service of one or two years rather than
3 Information published by Mr. Luis J. Reyes in the special forestry
edition of the Manila Daily Bulletin, summer of 1928.
2 Data furnished by the Bureau of Public Works, Manila.
189
190 The Philippine Journal of Science mi
submit to the inconvenience and expense of dismantling the
body for the purpose of replacing the timber. It is at best a
gamble as to the extent of renewals necessary, for a proper
estimate of cost can only be reached by fully exposing all the
wood. Estimates made on any other basis are usually high, in
order to take care of the probable deterioration of unseen parts.
The writer has rebuilt the bodies of two cars and in both cases
practically all of the wood needed replacement.
Often the deterioration becomes distinctly noticeable within
the first year's service, and there are many instances where ex-
tensive repairs have been necessary at the end of two years, or
even within one year. For cars in service longer than two years
it is safe to assume that decay is at least well started at some
important point (Plates 1 and 2). It impresses itself upon the
attention when the sills and vertical members have become suffi-
ciently decayed to permit the doors to sag and be thrown out of
alignment. Top members upon which any weight is hung, such
as the wind shield, also begin to rattle or give way entirely.
The decay may also spread to the upholstering or even affect
the top covering.
Repair work in the Philippines is not very thorough, as a rule,
partly as a result of the desire to keep the cost down, partly
through ignorance as to how wood decay develops and spreads.
Another factor in the situation is the trading in and resale of
used cars after conditioning them at the least possible expense.
A splice here and there, a few bolts tightened, a new coat of
paint, a new bit of upholstery, and perhaps a new top covering,
work wonders in appearance and often hide from the unwary the
more serious defects within. Even in what are considered
bona fide jobs the contractor often uses poor judgment in failing
to take out timber showing early stages of infection, little realiz-
ing that all traces of the wood-decaying organism must be eradi-
cated if the further spread of the decay is to be stopped. It is
often false economy to splice, and if the top framing or the sills
show considerable decay at the joints or elsewhere the better
procedure is complete replacement. This opinion is based on
the assumption that if the timber is so perishable as to rot out
at any point within a short period it will continue to do so, the
probabilities being that the replacements will out-last the orig-
inal timber left in and necessitate further repairs within a short
time.
46,2 Humphrey: Wood decay in Automobiles 191
WHAT CAUSES DECAY
Decay is conditioned upon moisture. Free water must gain
access to the wood in some manner. This may occur indirectly
by condensation when a cool surface is in contact with a satu-
rated atmosphere, or directly, through a leak in the top or else-
where, or when beating tropical rains force in moisture around
the closed doors and windows. Much wetting of the sills also
occurs through negligence in leaving doors or windows open
during storms, and some of it occurs during the process of fre-
quent washing.
Wet wood in a car dries but slowly, for it is usually covered.
When water reaches a joint it penetrates deeply and gets well
into the interior of the timber at the joined ends. This end
penetration of moisture into wood is very rapid and easily dem-
onstrated. It is thus a simple matter for water to enter at the
joints but very difficult for it to get out, even in rather dry
weather. When one stops to consider, however, that during a
tropical rainy season over 100 inches of rain may fall and weeks
may pass with the air at or near saturation, it is small wonder
that cars built of perishable wood deteriorate rapidly. Six
months under test conditions very highly favorable for decay
will destroy for practical use nearly all the temperate-zone woods
now used in American or European-made cars.
The next question that arises is what agent causes this dete-
rioration. It is all due to the presence of fungi belonging to
the more highly organized groups, principally the Hymenomy-
cetes. These fungi are plants fundamentally differing from
ordinary plants only in the lack of green coloring matter and
the method of nutrition. Ordinary plants must get their food
from the soil and air, therefore, they must have the green color-
ing matter to act as a catalyst in the manufacture of carbohy-
drates needed for growth. Fungi get all their food from the
organic substances upon which they grow, therefore, there is no
need for chlorophyll. They must, however, produce ferments to
render soluble and assimilable the various chemical substances
of which their substratum is composed. Wood-destroying fungi
are abundantly supplied with the ferments necessary to decom-
pose the compounds in wood, of which the principal ones are
cellulose and lignin, with some sugars and starches.
Wood-destroying fungi require for growth a small quantity,
of air, a favorable temperature, suitable kinds of wood, which
262412 3
192 The Philippine Journal of Science 1931
do not contain substances poisonous to, or inimical to, the growth
of the organisms, and a considerable amount of moisture. The
first and second requirements are met at all times in the Tropics,
where the temperature rarely goes below 60° F. and most of
the time is around 80°, or somewhat above. The third condition
is met when perishable woods such as are ordinarily employed in
automobile construction are used, and this is particularly true
of the sapwood of practically every species of tree known. The
third condition is readily met in the Tropics, where high humid-
ities and heavy rainfall prevail, and is accelerated by the,
factors mentioned in the third preceding paragraph. While
scientific knowledge of the exact amount of moisture most fa-
vorable to decay is lacking, enough data have accumulated to
indicate that the amount will vary for the kind of wood under
discussion. It is quite safe to say that ordinary absorption of
moisture from a saturated atmosphere (fiber saturation point)
is not sufficient for decay and that a certain amount of free
water must be present in the cavities of the wood cells or fibers
for the fungus to grow vigorously and break down the structure
rapidly.
LIFE CYCLE OP WOOD-DESTROYING FUNGI
The life cycle of wood-rotting fungi is quite simple. Each
fungus has two principal stages of growth, the sporophores, or
fruiting bodies, which take the place of the seed-bearing appa-
ratus of green plants, and the mycelium, which functions within
the wood as an absorbing system comparable to roots. The
mycelium is the stage that causes the damage. It consists
of fine cottonlike branched threads, which ramify through-
out the wood tissues and by the secretion of various ferments
cause their disintegration. These threads develop abundantly
in any closed-in moist space and may thus spread rapidly over
the surface of such inclosed timbers (Plate 2, figs. 7 and 9), as
well as within them.
After the wood becomes partially decayed the fungus attempts
to form fruiting bodies on the surface. These are very often
abortive and may consist only of cushions of compact mycelium
when developing in the dark, but when they have access to light
they take on a more or less definite form by means of which they
can be identified. Such fructifications developing in the light
soon become fertile; that is, the outer surface of such types as
grow in a thin layer flat against the wood, or the under surface
of shelving forms, produce large quantities of spores that are
46,2 Humphrey: Wood decay in Automobiles 193
comparable to seeds. These spores are very minute and easily
disseminated by air currents. When they lodge on the surface
or in the joints of moist wood, or even on wet cloth fabric, they
readily germinate to produce another crop or mycelial threads,
which in turn quickly penetrate and rot the material. In this
way organic construction materials are constantly subject to
infection, and when conditions are right for germination and
growth of the fungi, and the wood is not resistant to attack,
disaster comes.
Up to the present time only three species of fungi have been
observed fruiting on automobile wood; namely, Lenzites striata
(Plate 2, fig. 10), Polyporus sanguineus, and Trametes versa-
tilis, but there are, in all probability, a number of others that
have not been found in fruiting condition. These species are
inhabitants of warm regions and are well represented in the
Philippines as well as in the southern United States. They are
all species that are resistant to drying, to rather high temper-
ature, and to bright sunlight, as is evidenced by their frequent
occurrence on timber in the open, rather than in the shady, cool,
moist forest. There are a number of other such resistant
species that one would expect to find attacking automobile bodies,
exposed as they are to such fluctuating climatic conditions.
It appears probable that the infection of the wood occurs
after the cars reach the Tropics. This would certainly be the
case if all the wood going into their construction were kiln
dried before use, for the usual processes of artificial drying steri-
lize timber quite effectively.
PREVENTION OP DECAY
On this assumption then, what can be done by the manufac-
turer to adapt his product to tropical conditions? There appear
to be but two alternatives; either he must select the heartwood
of reputedly durable species of timber, or else the nondurable
woods now in use must be treated with a preservative.
It is said that durable tropical woods are being used in certain
European-made cars and that these are giving very good service
in the Tropics. American manufacturers may use a few exotic
woods for trim, but their inclusion as principal members has
never come to the attention of the writer. There are a great
number of Philippine woods that are well adapted to automobile
construction and that combine high resistance to decay with the
other mechanical properties desired. The difference in cost of
these superior tropical woods is a very small item in the total
194 The Philippine Journal of Science im
cost of manufacture and could readily be absorbed through the
increased business incident to the production of a car that would
stand up under severe conditions for a reasonable length of time.
It is absurd for a manufacturer to spend millions on improved
mechanical development and then house it all in a rotten shell.
Whether it would prove more economical to import these trop-
ical woods into the States for use in cars destined for the Tropics
or to thoroughly impregnate the nondurable American woods
now in current use is a matter for the manufacturer to decide
after checking up the respective costs. Either procedure would
be satisfactory from a durability standpoint. It is safe to say
that wood-frame cars in the Philippines are now at a serious dis-
advantage and are becoming increasingly unpopular. Steel is
giving good service and discriminating buyers are turning more
and more to it from the standpoint of both service and safety.
METHODS OF APPLYING WOOD PRESERVATIVES
The American manufacturer who wishes to use wood preserv-
atives has at his call a well-developed industry using standard
and proven processes, but if he should wish to treat his own
stock a suitable small pressure plant could be installed at a very
moderate cost.3 At present a small amount of treated timber
is being used in American-made cars, but it is insignificant and
not at all commensurate to the needs. It is unnecessary to go
into the details of treating methods other than to state that
pressure treatments are indicated. The principal commercial
substances injected into wood to increase its durability are coal-
tar creosote, zinc chloride, and sodium fluoride.
Creosote is the best all-around preservative known and is
particularly suitable for sills or timbers closed in by metal or
other impervious covering, for it is a brown to blackish oily
substance that would readily stain fabric. For other places a
colorless water-soluble substance like zinc chloride or sodium
ilouride is preferable, the former being used in a 6 per cent
concentration, the latter in a 3 per cent. Wood treated with
either of the latter substances can be satisfactorily painted if
necessary.
8 Full information can be secured from the Forest Products Laboratory,
Madison, Wisconsin, or from the American Wood Preservers' Associa-
tion, Chicago, Illinois.
46,2 Humphrey: Wood decay in Automobiles 195
It is also claimed 4 that creosoted wood can be satisfactorily
painted. The article cited states that "results thus far obtained
indicate that the use of aluminum paint on creosoted wood is
entirely satisfactory, providing the proper vehicle is used and
the wood is first allowed to dry for a time after treatment.' ' If
this be the case the sole objection to the use of creosote on ex-
posed parts of the automobile body would be overcome.
In replacement work preservatives could also be used to ad-
vantage in the Tropics. Most shops in Manila use untreated
guijo (Shorea guiso) for general repair work. This is a mod-
erately heavy to heavy wood, which is rather hard, tough, and
difficult to split. It is moderately durable and is widely used
in the Philippines for vehicle parts. Unfortunately, this wood
is refractory to preservative treatment. Apitong (Dipterocar-
pus spp., officially D. grandiflorus Blco.) is another Philippine
wood, however, that compares well in strength with American
white oak and that readily absorbs either creosote or water-
soluble preservatives, hence is widely used for treated ties, poles,
posts, piling, etc. There is no reason why it cannot satisfac-
torily be used in place of guijo, with the added advantage, when
properly treated, of being highly resistant to decay. There
should be no necessity for replacing treated apitong during the
life of the car. A proper treatment would be about 6 pounds
of creosote per cubic foot (empty cell pressure process), or ap-
proximately 0.5 pound (dry salt) of zinc chloride or 0.25 pound
of sodium fluoride.
As an alternative to this more-approved procedure, soaking
the timber, after framing, for several hours in a wood or iron
vat of the hot solution and then allowing it to cool in the same
solution to atmospheric temperature would also give a high
degree of protection, probably sufficient for the purpose in most
cases. Such a nonpressure process merely requires a vat of
suitable size fitted with steam coils to bring the temperature up
to the desired point. If creosote is used it should be heated to
about 180° C, while water solutions are brought to the boiling
point and held there until the wood is heated to the center. If
the vats are covered evaporation will not be excessive. The use
of iron vats would simplify heating where steam is not available,
since a fire beneath would accomplish the same purpose.
4 Wood Preserving News 8 (Dec. 1930) 177. Published by the American
Wood-Preservers' Association.
196 The Philippine Journal of Science
There is still another process in which the hot preservative
is applied as a spray or is put on with a wire-bound paint brush.
While this is hardly more than a make-shift it will increase the
life of the timber to some extent, particularly if the ends of the
timber, where joined, are allowed to absorb all the solution pos-
sible. This is better than no treatment at all, even with refrac-
tory wood such as guijo.
In Manila at the present time there is but one pressure treat-
ing plant,5 handling principally large construction material. It
is questionable, however, whether it would be feasible for the
small repair man to have his comparatively small quantities
of framed material treated on special order. Therefore, while
we do not usually recommend the cutting and shaping of timber
after treatment, it might be feasible, since apitong takes treat-
ment so readily, to have the blanks cut out to approximate size,
treated and held in stock either by retailers or by the plant.
There would be some wastage in such a procedure, but the mate-
rial cost in car-repair work is small compared to the labor cost.
Taking everything into consideration, however, the non-
pressure vat treatment, while less reliable when not properly
done, may be more convenient. If the treating equipment be
located at or near the place where the repairs are being made
it would be a simple matter to cut and fit the timbers for a
job, drop them in the preservative vat for a few hours, then
assemble them. Creosote-treated stock would not require more
than a couple of days in the sun to dry the surface, but of
course wood soaked in a water solution would require drying
again to its original air-dry condition. Small pieces would
season in a few days under cover during dry weather, but larger
pieces would probably require rather too long a delay for econo-
mic operation. In consequence of this it would seem advisable
to use creosote wherever possible.
9 Atlantic, Gulf, and Pacific Company.
ILLUSTRATIONS
Plate 1
Fig. 1. Body of 1925 model closed car removed for repair of the wood
frame. Note the severe decay in the sills and timbers joined to
them.
2. Decay of the upper right corner in the top of the same car.
3. Decay of running board after two years service in Manila.
Plate 2
Fig. 1. Thoroughly rotted frame of 1926 model closed car after three
years service in the Philippines. The timbers are so rotten
that the frame fell apart at the joints when the metal covering
was removed.
2. Front view of the same car.
3. Back left corner of the same.
4. View of back right corner and side of the same car.
5. Detail view of upper left corner at back (see fig. 3).
6. Left sill of same car.
7. Detail view of left corner (see fig. 3). Note fungous mycelium
clinging to the horizontal piece.
8. Sills of a 1921 model car decayed at the ends.
9. Fruiting bodies of Lenzites striata on decayed wood taken from
the frame of another closed car.
10. Sill from the car shown in Plate 1, fig. 1. The white coating
consists of a thin layer of fungous mycelium.
197
HUMPIlltKY : I>H'AY OK Wnoii.l
t'PlfllJF. .JuVHN. SCI., Uk N«l
^|vv#^^
PLATE 1.
Humphrey: Dkoay of Wooi>. |
F Philip. Jo^kx, Sri.. •!*>» Nn
PLATE 2.
COMPOSITION OF PHILIPPINE PEANUT OIL
By Aurelio 0. Cruz and Augustus P. West
Of the Bureau of Science, Manila
ONE PLATE
High-grade peanut oil serves as a salad oil and also for the
manufacture of oleomargarine. The lower-grade oil is suitable
for making soap. The oil cake from peanut oil serves as an
excellent cattle food as it contains a very high percentage of
proteins and is easily digested.1
Recently oil from Philippine peanuts was investigated in this
laboratory and the results showed that the Philippine oil has a
composition very similar to that of American peanut oil. It
would seem that there are promising prospects for the develop-
ment of peanut cultivation in the Philippines.
The composition of peanut oil has been the subject of a num-
ber of investigations as pointed out by Jamieson, Baughman,
and Brauns 2 in their paper on the composition of oil obtained
from the white Spanish type of peanuts grown in South Caro-
lina and also the Virginia type grown in Virginia. Their results
showed that the composition of the saturated acids obtained from
the glycerides of these two oils is about the same though the
Spanish type oil contains a slightly larger amount of saturated
glycerides than the Virginia type.
Information concerning the commercial aspects of the peanut
industry such as the picking and handling of peanuts, by-
products from crushing peanuts, and peanut oil, flour, butter,
candy, and cookies may be obtained from various Government
publications.3
Recently there have appeared two articles 4 which give a very
good resume of the present status of the peanut industry.
1 Lewkowitsch, J., Chemical Technology and Analysis of Oils, Fats, and
Waxes 2 (1922) 314.
2Journ. Am. Chem. Soc. 43 (1921) 1372.
3Beattie, W. R., U. S. Dept. Agr. Bur. Plant Ind. Cir. 88 (1911).
Reed, J. B., U. S. Dept. Agr. Bull. 1096 (1922).
Beattie, W. R„ U. S. Dept. Agr. Bur. Plant Ind. Cir. 98 (1912).
Bailey, H. S., and J. A. Le Clerc, Yearbook U. S. Dept. Agr. (1917) 289.
4 Lynch, D. P. J., Journ. Chem. Ed. 7 (1930) 794, 1037.
199
200 The Philippine Journal of Science imi
Peanuts are cultivated to some extent in the Philippines, but
to supply the local demand considerable quantities are also im-
ported.
During the year 1929, 1,632,960 kilograms of peanuts valued
at 256,833 pesos and 1,870,107 kilograms of peanut oil valued at
570,435 pesos were imported into the Philippines.5 Most of
these supplies come from China. Peanuts can be grown very
easily in the Philippines, both from cuttings and seeds, espe-
cially in rotation with rice, corn, and other short-maturing crops.
Since there is a considerable demand for peanuts and peanut oil
it would seem that their cultivation should offer excellent pros-
pects as a Philippine industry.
EXPERIMENTAL PROCEDURE
The peanuts used in this investigation were very kindly sup-
plied by Dr. Nemesio Mendiola, of the College of Agriculture,
University of the Philippines. They were the kind of peanuts
known locally as the Valencia variety. The shells were first
removed from the nuts after which the kernels were heated in
an oven (80° C.) for about an hour. As the heating expels
most of the moisture, the brown seed coats can then be removed
easily from the kernels. After removing the seed coats the ker-
nels were ground to a pulp, which was then cold-pressed to
obtain the peanut oil. The oil was filtered to eliminate most
of the fiber. After successive treatments (warming, shaking,
and filtering) with kieselguhr, suchar, and talcum powder, a
sample of oil was obtained with only a slight yellow color and a
very high degree of purity. The yield of oil, calculated on the
shelled nuts, was about 40 per cent.
The constants of Philippine peanut oil are given in Table 1.
There are also included for comparison the constants of Spanish
and Virginia type peanuts as determined by Jamieson, Baugh-
man, and Brauns.
Figures for the Philippine oil represent the average of closely
agreeing duplicate determinations. As shown by the data (Table
1) the physical and chemical constants of the Philippine oil are
quite similar to those of American oils.
The saturated and unsaturated acids that occur as glycerides
in Philippine peanut oil were separated by the lead-salt-ether
s Annual Report, Insular Collector of Customs, Manila (1930).
46,2 Crvz and West: Peanut Oil 201
Table 1. — Physical and chemical constants of peanut oil.
Constants.
Philippine oil
from Valencia
variety of
peanuts.
American oil.*
Spanish-type
peanuts.
Virginia-type
peanuts.
Specific gravity
0.9077,?^C.
4o
1.4676
b101.3
191.5
0.29
0.10
'17. 56
77.44
17.12
77.89
! 125.0
0.9148,?f^C.
25o
0.9136,ff-C.
25o
Refractive index
Iodine number
90.1
188.2
0.22
0.12
<*21.4
73.4
20.6
74.6
121.8
94.8
187.8
0.27
0.03
°17.4
77.7
16.4
78.7
118.2
Saponification value
Unsaponiflable matter per cent__
Acid value
Saturated acids, determined per cent__
Unsaturated acids, determined do
Saturated acids, corrected do
Unsaturated acids, corrected do
Iodine number of unsaturated acids, de-
termined
a Analyzed by Jamieson, Baughman, and Brauns.
b Determined by Hanus method.
c Iodine number 3.1.
d Iodine number 4.8.
• Iodine number 7.1.
method 6 in accordance with the suggestions of Baughman and
Jamieson.7 The results are recorded in Table 2.
Table 2. — Separation of saturated acids from the unsaturated acids in
peanut oil by the leadrsalt-ether method.
Experiment
No.
Oil used.
Unsatu-
rated
acids.
Saturated
acids.
Unsatu-
rated
acids (de-
termined).8
Saturated
acids (de-
termined).
Unsatu-
rated
acids (cor-
rected).
Saturated
acids (cor-
rected).
1
0.
9.5764
9.6772
0.
7.3966
7.5143
9-
1.7000
1.6805
Per cent.
77.24
77.65
Per cent.
b 17.75
o 17.37
Per cent.
77.68
78.09
Per cent.
17.31
16.93
2
Mean__
77.44
17.56
77.89
17.12
* Unsaturated acids: Saponification value 202.3; iodine number (Hanus) 125.0.
b Iodine number (Hanus) 3.10.
c Iodine number (Hanus) 3.15.
The composition of the unsaturated acids separated from
peanut oil by the lead-salt-ether method was determined by
means of the bromoderivative method.8 This consists in con-
verting the unsaturated acids into their bromo-derivatives, which
6 Lewkowitsch, J., Chemical Technology and Analysis of Oils, Fats, and
Waxes 1 (1921) 556.
7 Cotton Oil Press 6 (1922) 41. Journ. Am. Chem. Soc. 42 (1920) 2398.
8 Lewkowitsch, J., Chemical Technology and Analysis of Oils, Fats, and
Waxes 1 (1921) 585.
202 The Philippine Journal of Science 1931
are then separated by suitable solvents. The laboratory data
for duplicate analyses are given in Tables 3 and 4.
Table 3. — Determination of unsaturated acids of peanut oil (bromo-deriva-
tive method). Analysis 1.
Grams.
Sample of unsaturated acids 3.1512
Linolic tetrabromide insoluble in petroleum ether, melt-
ing point 113-114° C. 1.2013
Residue (dibromide and tetrabromide) ; bromine con-
tent, 40.37 per cent 4.3385
Dibromide in residue, 75.57 per cent 3.2786
Tetrabromide in residue, 24.43 per cent 1.0599
Total tetrabromide found 2.2612
Linolic acid equivalent to tetrabromide 1.0552
Oleic acid equivalent to dibromide 2.0918
Table 4. — Determination of unsaturated acids of peanut oil (bromo-deriva-
tive method) . Analysis 2.
Grams.
Sample of unsaturated acids 3.2970
Linolic tetrabromide insoluble in petroleum ether, melt-
ing point 113-114° C. 1.4786
Residue (dibromide and tetrabromide) ; bromine con-
tent, 39.60 per cent 4.2577
Dibromide in residue, 80.06 per cent 3.4087
Tetrabromide in residue, 19.94 per cent 0.8490
Total tetrabromide found 2.3276
Linolic acid equivalent to tetrabromide 1.0862
Oleic acid equivalent to dibromide 2.1748
A summary of these duplicate analyses (Tables 3 and 4) is
given in Table 5.
In Table 6 is given the composition of the mixed unsaturated
acids of Philippine peanut oil. There are also included the cal-
culated percentages of glycerides corresponding to these indivi-
dual unsaturated acids.
Saturated acids. — 'The saturated acids were separated from
Philippine peanut oil by the lead-salt-ether method and esterified
with methyl alcohol. The mixed acids were dissolved in methyl
alcohol and saturated with dry hydrogen chloride gas. The
mixture was then heated on a water bath (reflux) for fifteen
hours, after which it was treated with water and the ester layer
separated. The esters were dissolved in ether and the ethereal
**, 2 Cruz and West: Peanut Oil 203
Table 5. — Unsaturated acids of peanut oil; summary of analyses 1 and 2.
Acid.
Analysis.
Mean.
1
2
Linolic
Per cent.
33.49
66.38
Per cent.
32.95
65.96
Per cent.
33.22
66.17
Oleic
Total.. _
99 87
98.91
99.39
Table 6. — Unsaturated acids.
Acid.
Mixture of unsatu-
rated acids.
Glycerides
in original
oil.
Composi-
tion.
Propor-
tions in
original
oil.
Linolic .
Per cent.
33.22
66.17
Per cent.
25.88
51.54
Per cent.
27.04
53.86
Oleic
Total
99.39
77.42
80.90
solution washed with sodium carbonate solution and afterwards
with water. The ethereal solution was then dehydrated with
anhydrous sodium sulphate, filtered, and the ether removed by
distilling. The impure esters (83.29 grams) , which were yellow,
were distilled under diminished pressure. A preliminary dis-
tillation at about 3 millimeters pressure was made. The esters
(83.14 grams) were then redistilled at 3 millimeters pressure.
Data on the distillation of the esters are given in Tables 7 and 8.
Table 7. — First distillation of the methyl esters of the saturated acids;
pressure, 3 millimeters ; 83.29 grams of esters distilled.
Fraction.
Tempera-
ture.
Pressure,
Weight.
A
170-178
178-181
181-185
185-196
mm.
3
3
3
3
14.69
16.25
12.94
11.73
27.53
b :■
c ;
D. _ _ _
Total _ -
83.14
204
The Philippine Journal of Science
3 9S1
Table 8. — Second distillation of the methyl esters of the saturated acids;
pressure, 8 millimeters; 83 J4 grams of esters redistilled.
Fraction.
Tempera-
ture.
Pressure.
Weight.
From first distillation.
Second
dis-
tillation.
A and B
1
2
3
4
5
6
7
Residue __
°C.
170-173
173-176
176-186
186-203
203-225
225-230
230-238
mm.
3
3
3
3
3
3
3
9.
17.15
15.54
14.15
7.92
8.24
6.02
11.87
1.99
c - -
Do -
D _
Do --
Do --- --
Total --
82.88
TABLE 9.-
-Analyses of fractions obtained in the second distillation of the
mixed methyl esters*
Fraction.
1.
2.
3.
4.
5.
6.
7.
Iodine
number.
2.5
4.5
12.4
16.3
9.5
3.2
1.4
Saponi-
fication
value.
207.1
203.7
199.0
189.4
177.1
165.3
156.6
Mean
molecular
weight of
mixed
esters.
270.9
275.4
281.9
296.2
316.8
339.4
358.2
Composition of
mixed esters.
Satu-
rated.
Per cent.
97.90
96.22
89.58
86.30
92.02
97.31
98.82
Unsatu-
rated.
Per cent.
2.10
3.78
10.42
13.70
7.98
2.69
1.18
Mean
molecular
weight of
saturated
esters.
270.4
274.9
280.9
297.0
319.2
341.0
359.2
* Calculated iodine number of unsaturated methyl esters was 119. Calculated saponifi-
cation value of unsaturated methyl esters was 192.6.
Table 10. — Saturated acids corresponding to methyl esters in
each fraction.
Fraction.
Acids.
Palmitic.
Stearic.
Arachidic.
Lignoceric.
1
Per cent.
92.83
76.25
52.78
3.80
0.
15.92
11.85
7.47
0.30
Per cent.
9-
Per cent .
9>
Per cent.
9-
2
15.07
32.32
78.44
22.24
2.34
4.57
6.21
1.83
3
4
5
65.74
68.73
39.11
5.42
4.14
4.64
Q
24.56
55.84
1.48
6.63
1.92
7 - -
Residue*
Total
35.54
14.95
14.20
10.03
a Residue assumed to be methyl lignocerate.
46,2
Cruz and West: Peanut Oil
205
In Table 9, are given the analyses of fractions obtained in
the second distillation of methyl esters. From the data,
Table 9, there were calculated the amounts of the individual
acids corresponding to the methyl esters contained in the various
fractions. The results are recorded in Table 10 and were cal-
culated in accordance with the methods outlined by Baugh-
man and Jamieson in their investigations of Hubbard squash-
seed oil 9 and also American peanut oil.10
In Table 11, are given the composition of the mixed saturated
acids and the glycerides in the original sample of peanut oil
corresponding to these acids.
Table 11. — Saturated acids. a
Acid.
Mixture of saturated acids.
Glycerides
in original
oil.
Weight.
Composi-
tion.
Propor-
tions in
original
oil.
0.
35.54
14.95
14.20
10.03
Per cent.
47.57
20.01
19.00
13.42
Per cent.
8.14
3.43
3.25
2.30
Per cent.
8.54
3.58
3.38
2.38
Total _ --
74.72
100.00
17.12
17.88
1 When separated from peanut oil the corrected percentage of saturated acids was 17.12.
Table 12. — Composition of peanut oil.
Constituent.
Glycerides of:
Unsaturated acids-
Oleic
Linolic
Saturated acids —
Palmitic
Stearic
Arachidic
Lignoceric
Unsaponiflable matter,.
Total.
Philippine
peanuts
(Valencia
variety).
53.9
27.0
8.5
3.6
3.4
2.4
0.3
99.1
American peanuts.*
Spanish
type.
52.9
24.7
8.2
6.2
4.0
3.1
0.2
99.3
Virginia
type.
60.6
21.6
6.3
4.9
3.3
2,6
0.3
99.6
a The composition of the American peanut oil was determined by Jamieson, Baughman,
and Brauns, Journ. Am. Chem. Soc. 43 (1921) 1372.
9Journ. Am. Chem. Soc. 42 (1920) 156.
10 Journ. Am. Chem. Soc. 43 (1921) 1372.
206 The Philippine Journal of Science
The composition of Philippine peanut oil is given in Table
12. There are also included for comparison the analyses of oil
from the Spanish and Virginia types of peanuts.
The determined iodine number of Philippine peanut oil was
found to be 101.3 and the determined saponification value 191.5.
The calculated iodine number is 93.3 and the saponification value
188.4. The iodine and saponification values calculated from the
composition of the oil agree fairly well with the determined
values.
SUMMARY
The composition of Philippine peanut oil has been determined
and the results (Table 12) indicate that the Philippine oil has
a composition very similar to that of American peanut oil.
The percentage of linolic and palmitic glycerides is slightly
higher in the Philippine oil than in the American oils while the
percentage of the other glycerides is about the same or slightly
lower.
Peanuts can be grown easily in the Philippines. Since Phil-
ippine peanuts yield an oil of high quality and about the same
composition as American peanut oil, it would seem that there
are promising prospects for the development of peanut cultiva-
tion in the Philippines.
ILLUSTRATION
Plate 1. Peanuts growing at Agricultural College, Los Baiios, Laguna
Province, Luzon.
262412 — i 207
Ceuz and West; Peanut Oil J
I Philip. Journ. Scl, 46, No, 2,
/■■"
*~*l
\
\
■M
JV
\
PLATE 1. PHILIPPINE PEANUT PLANTS.
NEW OR INTERESTING ORIENTAL FERNS
By Edwin Bingham Copeland
Of the Herbarium, University of California, Berkeley
LYCOPODIUM EDAffOI Copel. sp. nov.
Phlegmaria laxa, caule deorsum fere 2 mm crasso, f oliis ibidem
tristichis remotis oblongis basi cuneatis, sursum gracile, foliis
ovatis brevissime petiolatis 6 mm longis 3.5 mm latis, acutis
haud acuminatis, basi rotundatis, coriaceis, viridibus; spicis
infra furcam inferam sporophyllis lanceolato-ovatis subacumi-
natis plerisque alternantibus sporangias duplo superantibus as-
persis, sursum sporophyllis triangulari-ovatis sporangias vix
vel paullo superantibus acutis interdum imbricantibus ; ramis
foliosis 10-12 mm, spicis 1.5 mm latis.
Palawan, Mount Mantalingajan, Bur. Sci. 77930 Edano,
April, 1929.
Among Philippine species, this is nearest to L. delbrueckii,
from which it differs in having ovate, instead of oblong, leaves,
much more slender spikes with short sporophylls except at the
base, and in the peculiar transition zone below the first dicho-
tomy of the inflorescence. Among the more broadly construed
species of the past, this might have been included in the L.
phlegmarioides of Baker; hardly, however, in that of Gaudi-
chaud, even if one ignores the description and figure of that
species as having shoots bilaterally, not radially, symmetrical,
with moderately dimorphous foliage leaves, as in Diphasium.
CYATHEA BONTOCENSIS Copel. sp. nov.
C. heterolobae affinis, trunco ignoto, stipite 20-25 cm longo,
rhachique gracilibus supra basin paullo crassiorem 3-4 mm
crassis, fulvis, minute asperulis, glabrescentibus ; fronde 85 cm
longa, 30 cm lata; pinnis infimis 6 cm longis, medialibus 20
cm longis, 5 cm latis, acuminatis, subsessilibus, rhachibus in-
feme praecipue deorsum paleis pallidis linearibus minute ciliatis
2-3 mm longis ornatis, apices versus paleis minoribus et pilis
etiam albidis sparse vestitis; pinnulis sessilibus, usque ad 28
mm longis, 6 mm latis, obtusis vel acutis, apud basin paullo
dilatatis et inciso-lobatis (infimis interdum ibidem pinnatis pin-
209
210 The Philippine Journal of Science mi
nula utroque latere una), alibi obscure serrulatis, costa inferne
deorsum squamulis minutis fulvis, sursum pilis ornata, lamina
glabra papyracea; venulis ca. 10-paribus, plerisque pinnatis;
soris inframedialibus, magnis, indusio brunneo, mox fisso, per-
sistente.
Luzon, Bontoc Subprovince, Vanoverbergh 8 IS, November-
December, 1910.
Nearly related to C. heteroloba, from which it is distinguished
by more slender and naked stipe and rachis, narrower and less
crowded pinules, and the presence of elongate paleae on the
minor rachises and of hairs on the costse. The two form an iso-
lated group.
CYATHEA CALOCOMA (Christ.) Copel.
A specimen collected by Fenix, Bur. Sci. 12711, at Sablong,
Benguet, is essentially identical with those from Mindoro. This
range would be commonplace in most genera, but is notable in
Cyathea. Possibly this species is more common than is sup-
posed, being unrecognized unless the base of the stipe is present.
CTATHEA CONTAMINANS (Wall.) Copel.
This is the commonest and largest of the Oriental tree-ferns,
the least exigent as to habitat, and the only species evidently
able to thrive in open places at moderate altitudes. It varies
much in ampleness, which may be due entirely to environmental
conditions. The largest specimens have just been brought in
by Ramos, Bur. Sci. 77170, from Cagayan Province. The pinna
is more than a meter long; the pinnules 15 cm, and the seg-
ments 4 mm wide, almost entire, and obtuse or only subacute.
In the other direction, several collections run below the size
typical of C. dementis.
Two large specimens collected by Ramos in 1912, in Cami-
guin de Misamis, a small volcanic island north of Mindanao,
are remarkable in other respects. No. 1U872 has the pinna 95
cm long, pinnules 17 cm long and 3 cm wide, and falcate, acute
segments, the sterile ones 4 mm wide, the fertile ones only 2.5
mm wide and separated by much more than their own width.
No. lhSUl has pinnules 15 cm long and up to 4 cm wide, the
fertile segments likewise narrow if merely serrate but more
closely placed and, therefore, more numerous ; but many of these
segments are dilated by the elongation of the teeth, to a width
of 5 mm, with the margin then fairly laciniate. The rachis
of the pinna is pale-tawny ; of the pinnule, dark-chestnut.
46,2 Cop eland: Oriental Ferns 211
CYATHEA DUPAXENSIS Copel. sp. nov.
Trunco ignoto ; stipite 30 cm longo, 1 cm crasso, atropurpureo,
spinis vix 1 mm longis multis horrido, deorsum superne paleis
atrocastaneis angustissimis 1-2 cm longis sparse vestito ; rhachi
deorsum castanea sursum fulva, glabra, inerme; pinnis infimis
valde reductis, medialibus 45-50 cm longis, 14-15 cm latis, sub-
sessilibus, acuminatis, rhachibus fulvis mox glabris sparsissime
muriculatis ; pinnulis maximis 8 cm longis (infimis paullo minor i-
bus), 12 mm latis, subsessilibus, caudato-acuminatis cauda ar-
gute serrata, etiam ad basin vix pinnatis, costa inferne sub-
glabra hinc inde squamulis minutis adspersa ; segmentis obliquis,
contiguis 6-7 mm longis, 3 mm latis, oblique subacutis ; obscure
serrulatis, pallide viridibus, papyraceis, ad costas inter soros
sparse squamuliferis alibi glabris; venis 7-8 paribus, inconspi-
cuis; soris stricte costularibus, 0.6 mm latis, exindusiatis vel
squamulis subtensis.
Luzon, Nueva Vizcaya Province, Dupax, Bur. Sci. 14291,
McGregor, March -April, 1912.
Distinguished from C. callosa by the absence of a developed
indusium; from C. caudata by smaller, thinner frond, more
densely spiny and less scaly base of stipe, and smaller sori.
Cyathea callosa and C. caudata are doubtfully different.
CYATHEA EDAftOI Copcl. sp. nov.
C. melanophlebise affinis, minor, trunco 1-2 m alto, 10-15 cm
crasso; stipite brevissimo, 1 cm crasso, rhachique atrocasta-
neis, muricatis spinis 0.5 mm longis sursum minoribus sparsis,
deorsum paleis linearibus castaneis 6 mm longis squamulisque
amorphis appressis sparse vestitis sursum glabrescentibus ;
fronde ovata, 45 cm lata, utrinque angustata, subtripinnata ; pin-
nis infimis 5 cm longis haud remotis, infimis fructiferis 10 cm
longis, medialibus 30 cm longis 11 cm latis, brevisti-pitulatis,
rhachibus inferne fuscis glabris vel primo ad insertiones pin-
nularum parce paleatis; pinnulis infimis reductis, medialibus 6
cm longis 18 mm latis, subsessilibus, breviacuminatis, costis
inferne nudis obscuris; segmentis infimis interse liberis (pin-
nulis secundariis) plerumque adnatis, sequentibus ala angusta
confluentibus, 2.5-3 mm latis, obtusis, obscure crenulatis, nudis
papyraceis, superne nigro-viridibus inferne olivaceis; venulis
ca. 8-paribus; soris costularibus, magnis, indusio praecipue
marginem versus aperto, denum persistente.
LUZON, Cagayan Province, summit of Mount Cagua, altitude
212 The Philippine Journal of Science 1931
1,300 meters, Bur. Sci. 78709 {type), and 78700 Edano, Octo-
ber-November, 1929.
This differs from C. melanophlebia Copel.,1 as both species
are known, in having decidedly smaller fronds, smaller spines
but rougher rachis, and in being still more free of palese on
any part of the frond; possibly it is a reduced mountain-top
form. Another nearly related species is C. halconensis. The
oldest species in the general group is C. caudata.
CYATHEA MERRILLII Copel. sp. nov.
Pseudohemitelia, trunco ignoto; stipite valido, atropurpureo,
spinoso spinis 2 mm longis minute castaneo-furfuraceo; rhachi
inerme, fusco-fulva, inferne glabra; pinna mediale 50-55 cm
longa, 18 cm lata, subsessile, rhachi fulva, glabra, sparsissime
muriculata; pinnulis medialibus 10 cm longis (infimis paullo
minoribus), 16 mm latis, caudato-acuminatis, subsessilibus, ad
basin pinnatis pinnulis n adnatis, alibi fere ad costam pinnatifidis,
costa inferne squamulis minutis plerisque angustis sparsis ves-
tita; segmentis haud contiguis, 10 mm longis, 2-3 mm latis,
acutis, integris, tenuiter papyraceis, superne obscure, inferne
laete viridibus, costis inferne squamulis minutis f ulvis plerisque
ovatis vestitis; venis 10-11-paribus, quarum ca. 8-paribus apud
costam furcatis et ibidem soriferis; soris vix 0.5 mm latis, in-
dusio ad squamam unilateralem basalem reducto.
Luzon, Benguet Subprovince, Merrill 7819, May, 1911.
Probably related to C. mearnsii of the same region, but thin-
ner, with more entire segments, and clearly distinguished by
the reduction of the indusium to a basal scale.
CYATHEA PUSTULOSA (Christ.) Copeland.
Cyathea pnstulosa (Christ.) Copeland, Philip. Journ. Sci. § C 4
(1909) 51.
Babuyan, Camiguin Volcano, Bur. Sci. 79615 Edano, March,
1930. Previously reported from Oshima and Formosa; new to
the Philippines.
CYATHEA SQUAMICOSTA Copel. sp. nov.
Rhachi valida, fulva, minutissime asperula setis minutis obsi-
ta et tuberculis parvis paucis sparsa; pinna mediale sessile,
50-55 cm longa, 17 cm lata, abrupte in apicem lanceolatam pin-
natam contracta, bipinnata, costa paleis fulvis sordidis lanceo-
latis 2 mm longis et ovatis vix 1 mm longis vestita, demum
glabrescente, muriculata; pinnulis maximis 9 cm longis, acu-
1 Philip. Journ. Sci. 38 (1929) 131.
46,2 Copeland: Oriental Ferns 213
minatis, sessilibus, basi 22 mm latis, ibidem pinnatis pinnulis "
sessilibus, alibi fere ad costam pinnatifidis, costa inferne
paleis deseriptis hie persistentibus vestita; segmentis conti-
guis vel imbrieatis, 3 mm latis, subfalcatis, apiee rotundatis,
integris, coriaceis, olivaeeis, eostis deorsum squamuliferis, aliter
glabris; venis ca. 10-paribus, immersis et inconspicuis ; soris
eostularibus, parvis ; indusio mox in cupulam brevem reducto.
Luzon, Benguet Subprovince, Mount Pauai, Bur. Sci. 8326
McGregor, June, 1909, altitude 2,100 meters.
This species is well marked by the minute roughness of
the rachis, the chaffiness, the very closely placed segments
with rounded apices, and the small, costular sori. It is from
a much-visited locality, and has been held twenty years for
description, in the hope that more complete material with
younger sori would be collected.
DRYOPTERIS CLEMENSIAE Copel. sp. nov.
D. gregis D. canescentis indusiis persistentibus, rhizomate re-
pente, lignoso, 2 mm crasso, paleis paucis parvis atrocastaneis
vestito, glabrescente ; stipitibus approximatis 8-15 cm longis,
versus basin squamulis paucis deciduis praeditis, alibi rhachibus-
que setis f ulvis curvatis vix 1 mm longis, dense vestitis ; f ronde
10-15 cm longa, 4-5 cm lata, basi truncata, acuminata, segmento
apicale deltoidea pinnatifida, alibi pinnata, atroviride, subcoria-
cea; pinnis utroque latere ca. 5, oppositis, sessilibus, lanceolato-
ovatis, infimis haud reductis sed subdeflexis et basi plus minus
angustatis, aliis basi cuneato-truncatis, acutis apice subfalcatis,
serrato-lobatis, eostis venisque minute pubescentibus ; venatione
irregulare, venulis extra seriem unam areolarum costalium pie-
risque liberis; soris sparsis ad venulas aut dorsalibus aut ad
anastomonoses impositis, indusiis parvis, nudis, orbiculari-reni-
formibus.
Luzon, Isabela Province, Mount Moises, M. S. Clemens 16490,
April, 1926. Type in Herb. Univ. Calif. 2851*86.
The irregular venation is suggestive of D. otaria, to which,
however, there is no near affinity.
DRYOPTERIS PARASITICA (L.) O. K.
LUZON, Cagayan Province, Pagikpik, Bur. Sci. 796U Edano,
on slopes in forest, altitude 1,000 feet.
Ferns bearing this name, or such predecessor-names as Ne-
phrodium molle, are well known in Philippine collections, but
this is the first known to me that fairly represents the species
as now construed. It is identical with several Formosan collec-
214 The Philippine Journal of Science 1931
tions, and like enough to those of southern China. The sori
are a single pair at the bases of most segments, a second pair
on some.
ATHYRIUM OPHIODONTUM Copel. ep. nov.
Diplazium caudice ignoto ; stipite alto, 1 cm crasso, basi atro-
castaneo paleis castaneis 2 cm longis basi 1 mm latis alibi angus-
tissimis remote et minute spinoso-dentatis membranaceis con-
tortis et intricatis vestito, sursum laete castaneo glabro, nullibi
muricato ; f ronde magna, tripinnatifida, apice acuminata pinnati-
fida ; pinnis medialibus 60 cm longis, 20 cm latis, caudato-acumi-
natis, stipitulis 5 cm longis protensis, rhachi potius sub lente
quam sub digite muriculatis ; pinnulis infimis quam sequentibus
minoribus longiusque (5 mm) stipitulatis, medialibus usque et
11 cm longis, acuminatissimis, basi 3 cm latis, profunde pin-
nati fidis sinubus rotundatis integris angustis, costa inf erne pa-
leis paucis minutis vestita superne angustissime alata, ala ad
basin venae quaeque interrupta et in dentem parvum protracta ;
segmentis oblongis, medialibus 10 mm longis, 3 mm latis, pleris-
que abrupte subfalcatis, tenuiter herbaceis, glabris, inciso-serra-
tis dentibus approximatis acutissimis rectis vel inflexis; venulis
hie simplicibus ca. 10-paribus (in pinnulis et segmentis inferio-
ribus saepe f urcatis et in dentes fissos protractis) ; soris costu-
laribus, brevibus demum confluentibus, indusio angusto, pallido.
Luzon, Cagayan Province, Peiiablanca, Bur. Sci. 77188 Ramos,
May 12, 1929 "in forest streams, at low altitude." Type in
Herbarium Bureau of Science.
This has the form and dissection of A. blumei, but the tex-
ture of the A. umbrosum group. Athyrium costulisorum and
A. tenuifolium are Philippine species of similar size and texture;
the former is fully tripinnate and the latter has muricate axes.
The narrow, crinkly, brown (not black) palese and the very
narrow and sharp teeth distinguish A. ophiodontum from all
its relatives.
ASPLENIUM FINLAYSONIANUM Wall.
Our specimen is A. macrophyllum Sw. Christensen construes
the name as having that application; and, likewise, seems cer-
tainly correct in construing A. integerrimum Hooker and Gre-
ville, Icones Filicum, Table 136, as the same species, in spite
of Hooker's own testimony, Icones Plantarum, Plate 937, that
it was an inaccurate presentation of the fern there (plate 937)
described as A. finlaysonianum Wall. The latter is based on
Wallich 2682, named A. hookerianum Wall, in the List — ac-
*6'2 Copeland: Oriental Ferns 215
cording to our copy and according to Christensen, Index, page
115. Although both Hooker, loc. cit., Plate 937, and Mettenius,
Asplenium, No. 149, cite Wallich 2682 as the type or basis of
A. finlaysonianum Wall., I mistrust their accuracy. It seems
to me that the proper name or citation for their plant, instead
of A. finlaysonianum Wall., or A. finlaysonianum Wall.; Hooker,
as in Christensen's Index, is A. finlaysonianum Hooker, non
Wallich.
The question then arises, could Hooker take Wallieh's name,
nomen nudum though it was, and, ascribing it to Wallich, give
it valid application to a different plant? Or did Wallieh's typi-
fication go with his specific name? If the latter is the case, the
A. finlaysonianum of Hooker has no valid name. In his Species
Filicum III, 272, published sixteen years after the plate in his
Icones, Hooker's first citation is Wallich 191, from Penang and
neighboring islands, as in the List, and his citation of No. 2682
is explicitly indirect. One might readily assume that No. 191
was a mixture, if it were not that all other published localities
(and all of our specimens) are Himalayan.
ASPLENIUM TRIPINNATIFIDUM Copcl. sp. nov.
Darea, ut videtur A. flaccido afiinis, caudice ignoto, stipite
longa rhachique straminee-viridibus superne profunde sulcatis
paleis minutis lanceolatis sparsis; fronde 75 cm longa, 25 cm
lata, acuminata, pallida, coriacea, inferne squamulis ovatis
0.4-0.7 mm longis ovatis clathratis castaneis sparsa tripinnati-
fida; pinnis infimis 8 cm longis, ovatis, medialibus 13 cm longis,
4.5 cm latis, valde caudatis, basibus stipitulatis oblique dila-
tatis, ad rhacheos applanatas vel anguste et crasse alatas pin-
natis; pinnulis inferioribus 3.5 cm longis, 5 mm latis, oblique
pinnatifidis, sequentibus (plerisque) linearibus incisis vel ser-
ratis, segmentis, resp. dentibus, 1 mm latis, acutis; venis om-
nino occultis; soris paucis, 2-3 mm longis, indusio 1 mm lato,
persistente, ad marginem non attingente.
Luzon, Rizal Province, Loher U379, April, 1913. Type in
Herb. Univ. Calif. 243202, distributed from the herbarium Bu-
reau of Science, Manila, as Tapeinidium pinnatum.
The texture, color, dissection, and squamules mark this as a
member of the chiefly Austral group of A. flaccidum and A.
bulbiferum. Taken in a broader sense the same group is rep-
resented by the widespread and common A. tenerum. It must
be construed still more broadly to make it include A. bullatum
Wall. ; Mett., which has glabrous fronds of different texture and
is very far from identical with A. bulbiferum.
216 The Philippine Journal of Science mi
STENOCHLAENA SMITHII (F&s) Underwood.
Lomariopsis smithii Fee, Acrostichum, p. 71, pi. 33 f. 2 and 53.
We have perfectly typical material of this species from Ca-
gayan Province, collected by Ramos in 1912, Bur. Sci. 18887.
Because it occurs here in typical form, I believe that it is rep-
resented also by Bur. Sci. 79645, collected by Edaiio at Pagik-
pik in the same province in 1930. This has the fertile pinna
(only one is present) nearly as broad as in typical S. smithii,
but the sterile pinnse quite like those of S. leptocarpa. The
rachis of the sterile frond is curiously wing-flattened toward
the apex. This mixture of characteristics makes me suspect the
distinctness of S. smithii and S. leptocarpa, neither of which is
represented in herbaria by enough material to establish its uni-
formity. Of the two names, S. leptocarpa has priority, due to
the apparent accident that Fee placed it in his section with
"frondibus homomorphis,,, although describing them as dimor-
phous.
LINDSAYA LONGA CopeL sp. nov.
L. gregis L. macraeanae, pinnulis profundius incisis, lamina
basiscopica imperfecte abscissa; rhizomate scandente, 2 mm
crasso, paleis late lanceolatis castaneis vestito; stipitibus alter-
nantibus castaneis, paleis paucis parvis ornatis, ca. 2 cm longis ;
fronde ca. 40 cm longa, 2.5-3 cm lata, utrinque attenuata, mem-
branacea, rhachi straminea; pinnis plerisque imbricatis, basi
acroscopice dilatatis, supra rhachin imbricatis, ibidem 6-7 mm
latis, deinde usque ad apicem plerumque acutam angustatis,
margine acroscopica prope basin i ad costam, apicem versus
profundius incisa, lamina basiscopica excisa, cum lobis infra api-
cem costam recipientam solitariis vel rarius nullis vel duo;
venulis in lobo quoque aut solitariis aut duo in soro anastomo-
santibus, alibi liberis; soris fere marginalibus, lunulatis basi
recurvatis, leviter si simplicibus, insigniter si venulas duas
recipientibus.
PALAWAN, Mount Balagbag, Bur. Sci. 77978 Edano, May, 1929.
The fronds are among the longest and narrowest in the group.
The pinnae are moderately protracted, less so than those of L.
apoensis and L. protracta, the other species having pinnse of the
same general form. The affinity to these long-stipitate species
is not as close as to the wide-spread L. macraeana and the
Philippine L. merrillii, from both of which L. longa is dis-
tinguished by the form and deeper incision of the pinnse and
the form and position of the sorus. It is quite distinct from
4*»2 Copeland: Oriental Ferns 217
L. fissa, likewise endemic in Palawan, which has more deeply-
cut pinnse with truncate lobes, and sori not at all lunulate.
PHILIPPINE OLEANDRA
Pedicel short and stout, less than 5 mm long and shorter than stipe.
Frond ciliate, pubescent.
Frond acute at base 0. mollis.
Base abruptly contracted 0. benguetensis.
Frond not ciliate, glabrous or sparingly hairy O. neriiformis.
Pedicel short, but stripe almost wanting 0. colubrina.
Pedicel commonly more than 5 mm long.
Frond sessile on the pedicel O. maquilingensis.
Frond stipitate.
Costa bearing long paleae 0. whitTieii.
Costa not conspicuously paleate.
Fronds firm, 30 cm or more long.
Sori 1 mm wide 0. cumingii.
Sori about 2 mm wide 0. macrocarpa.
Fronds smaller O. scandens.
OLEANDRA BENGUETENSIS Copel. sp. nov.
Rhizomate rampante, 4-5 mm crasso, radices graciles indi-
visas praelongas emittente, paleis 6-8 mm longis lineari-lan-
ceolatis integris vel subciliatis appressis nigrescentibus dense
obtecto; pedicello 2-3 mm alto, valido, sparse paleaceo; stipite
ca. 10 mm alto, costaque minute pubescentibus et sparsissime
paleaceis; fronde anguste lanceolata, ca. 25 cm longa, 2-2,5 cm
lata, acuminata, basim versus angustata basi ima truncata, mar-
gine angustissime cartilaginea minute ciliata, utraque facie
sparse puberula, papyracea; soris 2-6 mm a costa irregulariter
seriatis; indusio semiorbiculare sinu latissimo.
Luzon, Benguet Subprovince, Baguio, Elmer (Bureau of Gov-
ernment Laboratories) 6286, May, 1904. Type in Herb. Cope-
land 8225. Williams 1510, also from Baguio, September 24,
1904, probably represents the same species, but has a shorter
stipe, more paleaceous stipe and costa, and the sori closer to the
costa and in more regular lines.
Distinguished from 0. scandens by much stouter stems, with
supporting roots, shorter pedicel and stipe, longer, narrower,
less pubescent fronds, and indusia of different shape.
I have been abstaining from the publication of these (and
other) species of Oleandra, partly while I awaited publication
on Oleandra from Professor V. GoebeFs laboratory, partly in
the hope of learning what real 0. neriiformis is. As I now re-
cognize them, the published Philippine species are distinguish-
able by the above key, in which two or more species each are
included in 0. neriiformis and 0. colubrina.
218 The Philippine Journal of Science wi
OLEANDRA SCANDENS Copel. sp. nov.
Rhizomate repeote, 2-3 mm crasso, radices graciles enim fili-
formes ramosas multas emittente, ubique paleis persistentibus
castaneis lineari-lanceolatis 5 mm longis integris vel sparse et
irregulariter ciliatis supra basin peltatis dense vestito ; pedieellis
remotis vel subaggregatis, gracilibus, 10-20 mm altis; stipite
plerumque quam pedicello longiore, costaque pubescentibus ;
fronde lanceolata, 10-20 cm longa, 25-30 mm lata, acuta, basi
frondium minorum plerumque rotundata, ma jorum saepius
acuta, breviter ciliata, utraque facie pubescente, herbacea vel
subcoriacea; soris in lineam irregularem 2-5 mm a costa remo-
tam instructis ; indusio fere orbiculare versus costam sinu breve
angust affixo.
LUZON, Benguet Subprovince, Baguio, Elmer (Bureau of Gov-
ernment Laboratories) 6513, June, 1904. Type in Herb. Cope-
land 8219; Williams 1509, ibidem, August 1904; also from Ben-
guet, Copeland 180b, Bur. Sci. 2778 M earns, For. Bur. 15950
Bacani. Palawan, Silanga, Merrill 9850. While varying in
size, base, length of stipe and pedicel, and remoteness of
sori from the costa, these belong clearly to one species, well
marked by its pubescence, shape of indusium, and by fairly long
pedicel and stipe. I have chosen as the type of collection the
one probably most widely distributed to herbaria.
This has been distributed as O. cumingii Presl, a species de-
scribed with "frons fere sesquipedalis, coriacea," no mention of
ciliate border, which is too conspicuous and characteristic easily
to be overlooked, pubescence on the veins ; O. scandens is persist-
ently pubescent on lamina and veins.
Oleandra cumingii was based on Cuming 60 partim, as was
also O. mdcrocarpa Presl. My specimen of this collection
(sterile) can hardly be either of them, having a pedicel hardly
2 mm long. It is very hairy on surfaces and margin, and not
at all coriaceous. An imperfect specimen in the herbarium
University of California conforms perfectly to the diagnosis.
To O. cumingii has been reduced 0. chinensis Hance, described
as valde coriacea.
GRAMMITIS LIMAPES Copel. sp. nov.
Species G. pubinerviae et G. (Polypodio) bulbotrichae affi-
nis rubustior; rhizomate repente vel suberecto, 1.5-2 mm crasso,
paleis ferrugineis ovatis 1-2 mm longis vestito; stipitibus ap-
proximate, basi articulatis, 5-7 cm altis, 1 mm crassis, pilis
1 mm longis atropurpureis debilibus vestitis et ob baseos bulbo-
46,2 Cop eland: Oriental Ferns
219
sas pilorum omnino horridis; fronde vulgo 15, rarius usque ad
30 cm alta, maxima 9 mm lata, utrinque sed ad basim imam
abrupte angustata, integra, inferne praecipue ad costam deor-
sum setis minutis sparsissimis deciduis vestita aliter glabra,
rigide eoriacea, costa valida inferne prominente nigra; venis
furcatis et, ubi satis lata frons, ramo basiscopico item furcato,
ramo acroscopico breve; soris medialibus vel cost® paullo pro-
pioribus, fere superficialibus, 2 mm latis, 2-3 mm longis haud
confluentibus, sporangiis setis nigris brevibus obsitis.
Java, Gedeh, Panggrango, Copeland, May, 1915 altitude 2,800
meters.
This differs from Gedeh specimens identified as G. pubinervia
m being decidedly stouter and more rigid, with larger sori, as
well as in the peculiarity of the stipes, exceedingly rough to
the eye and to the touch. Grammitis pubinervia and G. con-
gener, as I construe them, have ordinary hairs on the stipe,
stiffer and rather longer than those of G. limapes, but without
enlarged bases. The Philippine G. bulbotricha, Polypodium bul-
botrichum CopeL, Philip. Journ. Sci. 40 (1929) 309, is likewise
less robust and with smaller, as well as more costal, sori.
GRAMMITIS MULTIPOLIA CopeL sp. noy.
Species jamdiu confusa G. pusillae Blume (Polypodio hirtello)
affinis frondibus longe stipitatis distincta; caudice breve, erecto,
parvo, basibus longe stipitatis distincta; caudice breve, erecto,
parvo, basibus stipitum radicumque occulto, paleis etiam occultis
lanceolato ovatis castaneis integris acutis vix 1 mm longis vestito;
stipitibus confertissimis, filiformibus, fuscis, 2-3 cm longis, pilis
atropurpureis usque ad 2.2 mm (plerisque ca. 1.2 mm) longis
ornatis; fronde lineare, vulgo 6 cm rarius usque ad 9 cm longa,
4 mm lata, obtusa, deorsum sensim angustata, integra vel sub-
integra, praecipue inferne sparse setosa, subcoriacea, costa
gracile inferne prominente; venis immersis, furcatis, ramo
acroscopico vix basiscopico aequente et deorsum sorifero; soris
parvis, superficialibus, fere orbicularibus, subcostalibus, spo-
rangiis interdum setiferis.
JAVA, Mount Panggrango, Copeland, May, 1915 altitude 2,700
meters; ibidem, Miller, 1897.
I believe this to be Polypodium alpestre Blume non Spenn.,
subsequently regarded by Blume as a variety of his Grammitis
pusilla, but ill depicted in Flora Javae II, PL 46, f. 5, which
shows fronds too broad and too hairy. It differs from the typi-
cal form of that species in having long stipes, relatively nar-
220 The Philippine Journal of Science 1931
rower and firmer fronds and comparatively remote sori. It
seems to me decidedly more distinct than Blume's var. f3 lasio-
sora, construed as a species by Fee (Grammitis nana) and
Hooker.
GRAMMITIS STENOCRYPTA Cope!, sp. nov.
G. f asciatae similis, paleis angustis et soris elongatis prof unde
immersis costa remotis facile distinguenda, rhizomate brevire-
pente, 1.5 mm crasso, paleis ferrugineis 4 mm longis, 1 mm latis
dense vestito; stipitibus approximatis, 6-7 cm longis, 0.6-0.7 mm
crassis, pilis paucis debilibus deciduis aspersis, basi nigro bul-
bosa paleis rhizomatis immersa articulatis; fronde lineare, 20
cm longa, 7-8 mm lata, utrinque attenuata sed apice ipsa ob-
tusiuscula, integra, coriacea, inferne preaecipue ad costam pilis
obscuris vix 1 mm longis mox omnibus caducis vestita deinde
glabra; costa manifesta vix prominente; venis plerisque fur-
catis, ramo inferiore rarius iterum f urcato, ramis in hydathodis
inconspicuis terminantibus ; soris inframedialibus, prima ap-
paritione in cryptis 2.5-3, rarius usque ad 4, mm longis linea-
ribus immersis, deinde sporangiis evolutis ellipticis oblique po-
sitis, sporangiis setis brevibus protectis.
Java, Gedeh, prope Kandang Badak, Copeland, May, 1915.
At first sight, much like G. fasciata, and perhaps in the past
confused with that species. Too naked for confusion with G.
setosa or G. pubinervia, longer stalked with longer sori, farther
from the costa. Grammitis longa Fee can hardly be G. fasciata,
as it has been construed, because G. fasciata is remarkable for
its naked fronds and sporangia. The description of G. longa,
6th Memoire, page 6, Plate 4, f . 1., is far inferior to its author's
usual standard; it appears to differ from G. stenocrypta by
having short stipes, more divergent and mostly twice forked
sterile veinlets and the sori shorter, more costal, and more nearly
parallel to the costa.
CAMPIUM SUBSIMPLEX (Fee) Copel.
Acrostichum zollingeri Kze., Bot. Zeit. 4 (1846) 419.
The suspicion that these are identical, expressed in Philip.
Journ. Sci. 37 (1928) 357, can now be confirmed, as the Cali-
fornia Herbarium has come into possession of a sheet of Zol-
linger 1293. Among the names borne by this sheet is Lepto-
chilus lanceolatus.
EFFORTS TOWARD BIOLOGICAL CONTROL OF THE
COMMON PINK MEALYBUG TRIONYMUS
SACCHARI (COCKERELL) OF SUGAR
CANE ON NEGROS
By F. C. Hadden
Assistant Entomologist, Experiment Station of the
Hatvaiian Sugar Planters' Association
and
A. W. Lopez
Entomologist, Philippine Sugar Association
While the pink mealybug of cane is not of as much impor-
tance on Negros as it is on Luzon, still it is of sufficient impor-
tance to warrant attempts at more complete control. The insect
is numerous at present, perhaps due to the unusually dry rainy
season just experienced on Negros. The effect of the dry period
is to inhibit the growth of the entomophagous fungus Asper-
gillus sp., which is of considerable importance in the natural
control of the mealybug. The damage caused by mealybugs is
occasioned by the fact that they extract the cane sap and reduce
purities.
In order to try to decrease the number of mealybugs now in
evidence and to try to provide insurance against any possible
future outbreak of the pest, two kinds of natural enemies (from
Laguna, Luzon) have been liberated in parts of Negros by the
Entomology Department, Philippine Sugar Association.
One of the insects is Scymnus sp.1 (order Coleoptera, family
Coccinellidse) a small brown lady-bird beetle measuring 1.5
millimeters in length. The fully grown larvae are only 3 milli-
meters in length, their small size enabling them to get down
between the leaf -sheath and stalk where the mealybug is most
commonly found. They devour the young mealybugs, and are
thus predators.
3 According to a recent letter from Mr. Swezey this coccinellid is a species
of the genus Pullus.
221
222 The Philippine Journal of Science wn
The senior author believes that he is the first to discover the
Scymnus and its importance. The life history is completed in
about a month.
The second natural enemy is a small wasp, Anagyrus sp.
(order Hymenoptera, family Encyrtidae), determined by Mr. O.
H. Swezey, entomologist of the Hawaiian Sugar Planters' As-
sociation Experiment Station, from specimens sent to him. In
Hawaii there is another species of encyrtid that perfectly con-
trols the gray sugar-cane mealybug, but never attacks the pink
mealybug. It is very difficult to find gray mealybugs in Hawaii
to-day, but at one time they were so numerous that they were
considered destructive. This control was effected only after
the encyrtid became established in Hawaii. Efforts are now
being made to establish this new species of Anagyrus in Hawaii
and it is hoped that it will do as good work on the pink mealy-
bug as the other encyrtid does on the gray mealybug.
This family of wasps lives as parasites of the ova, larvae,
or pupae of various insects. In the present case, its eggs are
laid in nearly mature or mature mealybugs, the larvae probably
devouring the entire body contents. The length of the female
wasp, which is yellowish, is about 1 millimeter. The males,
which are black, are noticeably smaller. The life history of the
Philippine Anagyrus is from twelve to sixteen days depending
on the temperature. This wasp was first reared at the Col-
lege of Agriculture by one of the entomology students.
The senior author, temporarily stationed at the College of
Agriculture making studies of mealybug natural enemies for
the Hawaiian Sugar Planters' Association Experiment Station,
devised methods of rearing the two species above mentioned.
He advised the junior author of these methods and supplied him
with a supply for establishment on Negros.
Starting with a small nucleus, Mr. F. P. Goseco, assistant
entomologist, Philippine Sugar Association, has been able to
rear large numbers of both kinds of natural enemies in the labor-
atory. They are reared in cloth-covered battery jars and are
supplied with mealybugs as needed.
Up to December 23, 1930, colonies ranging in number from
forty to one hundred individuals of each species have been liber-
ated at the La Carlota Central Experiment Station field; at
the Ma-ao Central parent field ; at Hacienda San Jose, of Ramon
Yusay, Binalbagan Estate; at Hacienda Panaquiao of Emilio
Montilla, Isabela Sugar Company; at Hacienda Tarog of Ilde-
46,2 Hodden and Lopez: Control of Pink Mealybug 223
fonso Doronila, Santos-Lopez Central, Panay; at the Hawaiian-
Philippine Company experimental field ; and at the experimental
field of the Bacolod-Murcia Milling Company.
Fields are selected in which mealybugs are plentiful, and
which will not be harvested before January, 1931, thus allowing
the natural enemies sufficient time to become established.
Stocks are maintained in the laboratory of the Philippine
Sugar Association at La Carlota Central and further liberations
will be made in the near future.
262412 5
THE KAHN TEST IN CLINICAL SYPHILIS 1
By Carlos Monserrat
Of the Department of Pathology and Bacteriology, College of Medicine
University of the Philippines, Manila
Since the discovery of the Kahn test, numerous papers dis-
cussing the correlation between this test and the Wassermann
reaction have been published. However, in the majority of
cases, the problem has been approached too much from the serol-
ogical side of the question. The results are generally based on
the total number of cases examined, without much attention
being given to the presence or absence of clinical manifestations
of syphilis. Comparatively few studies have been submitted
in which the serological diagnosis was accompanied by clinical
data, especially in regard to the type of syphilitic lesions. In
the majority of these reports the percentage of agreement and
disagreement is also based on the total number of cases ex-
amined, including a large number of nonsyphilitic cases.
It is the purpose of this paper to add something to the gen-
eral knowledge of the Kahn reaction and to judge the merits of
the test in comparison with the water-bath and ice-box Wasser-
mann fixation methods, as performed in the serological labor-
atory of Johns Hopkins Hospital. The author tried to place
himself in the position of a clinician, who first makes the clinical
diagnosis and then receives further information from the serol-
ogical laboratory.
In these records, the writer has endeavored to demonstrate the
relative sensitiveness of the Wassermann and the Kahn tests
by means of a series of syphilitic cases only, corroborated in
each case by the clinical history of the patient.
Parallel Wassermann and Kahn tests were also made with
a smaller number of cases showing different clinical types of sy-
philis that had been submitted to several courses of treatment.
The object here was to get an idea of the merits of these reac-
tions as a guide in the treatment of syphilis.
lrThis paper is the result of work done in the School of Hygiene and
Public Health, Johns Hopkins University, Baltimore.
225
226 The Philippine Journal of Science issi
The clinical material consisted of syphilitic cases entering the
Department of Syphilology in the dispensary of Johns Hopkins
Hospital. The majority of these cases were selected and diag-
nosed by Dr. H. Hopkins, of the clinical staff of the hospital.
Only a few of the unselected cases submitted to the serological
laboratory are included.
The technic followed in the Kahn test is the procedure de-
scribed by Kahn in the latest edition of his book2 in which
incubation is almost entirely eliminated and readings are made
shortly after mixing the serum and the antigen. In the original
method, the weaker reactions required an overnight incubation
before the final reading was made.
The antigen used in the experiments here reported was pre-
pared with great care and titrated against a standard antigen
obtained from Doctor Kahn's laboratory.
Finally, the comparative tests were made upon the sera from
twenty-four hours to four days after bleeding the patients.
Under these circumstances, it was found that the age of the
serum did not interfere with the results of the precipitation test
if the material was kept properly in the ice box.
The Wassermann reaction was performed in the laboratory of
Johns Hopkins Hospital.3 A 0.2 per cent cholesterinized beef-
heart antigen was used, and the fixation carried out both in the
water bath and ice box. In the water-bath method, a first in-
cubation (water bath) of thirty minutes was allowed for fixa-
tion. In the ice-box method, this first incubation was carried out
in the refrigerator for three hours. After the first incubation
the hemolytic system was added ; for all the tubes were replaced
in the water bath at 37° C. for one-half hour before the readings
were made.
It will be seen from Table 1 that the cases have been classified
in seven groups, as follows :
1. Primary syphilis. Cases with early or more or less healing chancres.
2. Secondary syphilis. Cases with different varieties of skin and mu-
cous membrane lesions.
3. Tertiary syphilis. Skeletal, visceral, and late cutaneous involve-
ments.
2 Serum Diagnosis of Syphilis by Precipitation; governing principles,
procedure, and clinical application of the Kahn precipitation test. Wil-
liams and Wilkins Co. (1925).
8 A complete description of the method is given in an article by Albert
Keidel and Joseph D. Moore, The Wassermann reaction in the Johns
Hopkins Hospital, Johns Hopkins Bull. 34 (January, 1923) 16.
46,2
Monserrat: Kahn Test in Syphilis
227
4. Latent syphilis. Cases in which the Wassermann reaction has been
found repeatedly positive. The majority of these cases gave his-
tory of chancres, secondaries, or other manifestations and at the
time when the blood examination was made were inactive.
5. Syphilis of the central nervous system. Cases of neuro-recurrens,
general paresis, tabes, asymptomatic, and other nonspecified le-
sions of the central nervous system.
6. Congenital syphilis.
7. Nonsyphilitic. Cases with various skin diseases and other clinical
manifestations (impetigo, pytiriasis rosea, acne, eczema, derma-
titis seborrheicum, carcinoma, pregnancy, abscesses, arteriosclero-
sis, colloid goiter, fractures, chronic myeloid leukemia, psycho-
neurosis, endocarditis, and pericarditis).
Table 1. — Showing the number of cases examined and the various types of
the disease.
Type of syphilis.
Number
of
cases.
Remarks.
Primary
11
Syphilitic chancres. Some positive for treponema.
("Skin lesions, 34 cases.
Secondary
54
ISkin and mucous membranes, 11 cases.
Early skin with chancre, 4 cases.
'•Mucous membrane lesions, 5 cases.
f Skeletal lesions, 9 cases.
Tertiary
29
\ Visceral (vascular, cardio-vascular, eye) 11 cases.
[Skin late lesions, 9 cases.
Latent syphilis
35
/Neuro-recurrens, 1 case.
Central nervous system...
21
General paresis, 4 cases.
1 Tabes, 5 cases.
1 Asymptomatic and not specified, 11 cases.
Congenital syphilis
2
Nonsyphilitic
68
Total _._
220
The number examined was one hundred fifty-two syphilitic and
sixty-eight nonsyphilitic cases. It must be noted, however, that
one hundred forty-one cases of the syphilitic series had received
treatment before the time the Wassermann and the Kahn tests
were performed. In the syphilitic series there were only eleven
untreated cases with secondary lesions, and the serological re-
sults in these cases showed complete agreement, as demonstrated
in Table 5a. No false results were found in the sixty-eight
nonsyphilitic cases, and the results of the three reactions also
showed complete agreement.
Primary syphilis. — Table 2 shows that in eleven cases of pri-
mary syphilis the Kahn test gives more definitely positive
reactions than the Wassermann water-bath and ice-box fixation
methods.
228
The Philippine Journal of Science
1931
Table 2. — Showing the results of the Wassermann and Kahn tests in ele-
ven cases of primary syphilis.
[Very strongly positive, 4~+++ ; strongly positive, ■[' 14' ; moderately positive, '44- J slightly
positive, 4~ I doubtfully positive, + ; negative, — .]
Number of cases.
Kahn.
Wassermann.
Water bath.
Ice box.
+ + + +
+ + + +
4-4- + +
+ + + +
+ + +
+ + +
+
+ + + +
±
+ + + +
+ + + +
+ + + +
+ +
+ J-11313 (1-26-29).
JJ-15556 (11-19-29).
lJ-15121 (11-18-29).
2 __ _
1
3 _
Case J-11313. November 21, 1928. Patient had a ragged crusted ulcer
on shaft and a soft dirty shallow ulcer in coronal sulcus. Noticed sore in
penis November 3, 1928; exposure six weeks prior. Sore grew worse until
present. Dark field, negative. Wassermann 4-4, November 21, 1928.
Clinical diagnosis: Syphilis, primary. Seropositive.
Table 3. — Showing the record of treatment and reactions of
cases J -113 IS.
Date.
Treatment.
Dose.
Wassermann.
Kahn.
W. B.
I.B.
November 21, 1928
Neoarsphenamine
do
a.
0.60
0.90
0.40
0.40
0.40
+ + + +
+ + + +
+ + + +
+++ +
++++
++++
+
+ + +
December 1, 1928
January 11, 1929
do_ _._
January 18, 1929
do .
January 26, 1929 .__
do
Case J-15556. Patient came to the hospital January 26, 1929, with an
ulcer 0.5 by 0.5 centimeter on foreskin near corona. It was covered with
a dirty yellow slough and was very tender. Had been treated locally with
calmin and peroxide. Dark field, negative. Wassermann, negative (Jan-
uary 26, 1929). No treatment. February 4, 1929, Wassermann = Neg.
Neg. February 19, 1929, the penile lesion had a characteristic rolled, in-
durated border of the syphilitic chancre. Dark field, positive for trepo-
nemas. Wassermann Neg. Neg. Kahn 3 (February 19, 1929). One in-
jection of arsphenamine given February 19, 1929. February 26, 1929,
Wassermann = Neg. Neg. Kahn 3. Diagnosis: Primary syphilis.
Case J-15121. January 18, 1929, a subacute gonococcus infection. The
urethral orifice eroded by a superficial dime-sized soft ulcer. Bilateral
4G, 2
Monserrat: Kahn Test in Syphilis
229
small, hard, indolent, inguinal buboes. Dark field from ulcers showed many
nonmotile treponemas. Wassermann reaction = Neg. (water bath) = 4
(ice box). Clinical diagnosis: Syphilis primary. Seropositive.
Table 4. — Showing the record of treatment and reactions of case J -15121*
Date.
Treatment.
Dose.
Wassermann.
Kahn.
W.B.
I.B.
January 18, 1929
Neoarsphen amine....
do
0.90
0.90
0.90
January 25, 1929
—
+
+ + +
February 18, 1929
do
Secondary syphilis. — In the forty-three cases of secondary sy-
philis, the Kahn test was found also more sensitive than the two
Wassermann methods employed. See Table 5a.
Table 5a. — Showing the results of the Wassermann and Kahn tests in\
forty -three cases of secondary syphilis.
[Very strongly positive, ■ I ■ I ■ 1 "— h- ; strongly positive, +-H- ; moderately positive, ++ ; slightly
positive, +; doubtfully positive, ±; negative, — .]
Number of cases.
Kahn.
Treated cases.
Water bath.
Ice box.
16
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ +
+ + + +
+ +
+ + + +
2
+ + + +
2
+ + + 4-
2__
+ -f- + + J-16015 (V-3-29).
1__ ___
+ + +
1__
+ +
7 _.
fJ-11391 (111-13-29).
tj-16247 (IV-10-29).
— U-7696 (111-20-29).
1_
1___ __
11
UNTREATED CASES
11
+ + + +
+ + + +
+ + + +
Case J-16015. February 12, 1929, complaint of pain in the joints. Ker-
nels behind ears. Examination showed three large mucous patches on
hard and soft palate. Treponemas were demonstrated in these lesions.
Post auricular glands very large and firm. Old scars on prepuce. Palms
and soles showed dark, firm papules. Clinical diagnosis: Secondary cuta-
neous palmar and plantar. Arthralgia.
230 The Philippine Journal of Science 1931
Table 56. — Showing the record of treatment and reactions of case J -1601 5.
February 12, 1929 _
February 20, 1929.
February 27, 1929.
March 6, 1929
March 13, 1929.. _
March 20, 1929...
April 5, 1929
April 12, 1929
April 19, 1929
April 26, 1929
May 3, 1929
Treatment.
Arsphenamine.
do
.do.
-do.
.do.
.do.
_do_
_do_
Bismuth.
do...
do-
Dose.
G>
0.60
0.60
0.60
0.60
0.40
0.40
0.40
0.40
0.20
0.20
0.20
Wassermann.
W.B. I. B
+ + + +
+ + + +
+ + + +
+ + + +
+ + +
+ + + +
Kahn.
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ +
+ + + +
+ + + +
Case J-11391. January 14, 1929, numerous follicular indurated papules
over bearded region of face, some with tendency to be annular. No lesions
on mucous membrane. Penis distorted by large multiple scars and crusted
ulcers. Dark field from one of these, negative. Wassermann 4-4 (Jan-
uary 14, 1929). Clinical diagnosis: Secondary early cutaneous folliculo-
papular.
Table 6. — Showing the record of treatment and reactions of case J-11391.
Date.
January 15, 1929. _
January 22, 1929. _
January 29, 1929..
February 6, 1929. _
February 12, 1929.
February 19, 1929.
February 26, 1929.
March 5, 1929
March 12, 1929 ...
March 13,1929...
Treatment.
Neoarsphenamine..
do„
.do.
.do..
_do_.
.do.
.do.
.do.
Bismuth..
do...
Dose.
0.90
0.90
0.75
0.75
0.75
0.75
0.75
0.75
0.20
0.20
Wassermann.
W. B. I. B
+ + + +
+ + + +
+ +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
Kahn.
+ + + +
+ + + +
+ + + +
Case J-16247. February 13, 1929, a condylomata on right inner thigh,
fading papular lesions (about two to three months duration) over arms,
palmar and plantar macules. Patient stated that her husband had similar
eruptions before she had and was receiving treatment. Wassermann =
4-4. Clinical diagnosis: Secondary early pigmentary condylomata.
46>2 Monserrat: Kahn Test in Syphilis 231
Table 7. — Showing the record of treatment and reactions of J-162&7.
Date.
Treatment.
Dose.
Wasseimann.
Kahn.
W. B.
LB.
February 13, 1929
Neoarsphenamine
do
0.75
0.75
0.75
0.75
0.75
0.75
0.75
0.20
0.20
+ + + +
+ + + +
February 20, 1929
February 27, 1929
do___
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + +
March 6, 1929
do
March 13, 1929_
do
March 20, 1929
do
April 4, 1929
do
April 10, 1929
Bismuth
April 19, 1929
do
Case U-7696. January 30, 1929. Complained of soreness on genita-
lia. Several eroded condylomata lata on vulva were found. Treponemas
found by dark field. Wassermann = 2-4. Clinical diagnosis : Secondary
early condylomata.
Table 8. — Showing the record of treatment and reactions of case U-7696,
Date.
Treatment.
Dose.
Wassermann.
Kahn.
W.B.
I.B.
January 30, 1929
Neoarsphenamine
do _
0.75
0.75
0.75
0.75
0.75
0.75
0.75
0.75
0.20
February 6. 1929
+
+ +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
-4-
February 13, 1929
do
February 20, 1929
do
February 27, 1929
do_._
+
+ + + +
+ +
March 6, 1929__
do
March 13, 1929
do
March 20, 1929 _
do.
March 27, 1929..
Bismuth
Tertiary syphilis. — In the twenty-nine cases of tertiary syphi-
lis, as shown in Table 9, the Kahn test was much more sensitive
than the Wassermann reaction, especially the water-bath method.
In the late cutaneous involvements, the sera examined agree al-
most completely with the three methods. In the skeletal and vis-
ceral syphilis, the Kahn test is definitely more sensitive than the
water-bath method. See Table 9.
232
The Philippine Journal of Science
1931
Table 9. — Showing the results of the Wassermann and Kahn tests in
twenty-nine cases of tertiary syphilis^
[Very strongly positive, ++-++ 1 strongly positive,
moderately positive, +-+; slightly
positive, +; doubtfully positive, ±; negative, — .}
Type.
Number
oi
cases.
Kahn.
Wassermann.
Water bath.
Ice box.
Skeletal (9 cases) •
3
1
1
1
1
2
2
2
1
1
1
1
1
1
1
3
1
1
1
3
+ + + +
4-4-4-4-
4-4-4-
+ + + +
+ + + +
4.4-4-4.
+ + + +
+ + + +
4-4-4-4.
+ + + +
4.4-4-
+ +
+ + + +
4-4-4.4-
4-4.4-4-
+
+ + + +
+
4.4-4.4-
4-4-4-4-
4-4-4.
+ +
4-4.4.4.
4-4.4-4.
+ + + +
+ + J-12727.
— (G-83908 (1-23-1929).
4-4.4.4.
4.4-4-4-
+ + + +
+ +
— No. 20149 (1-10-1929).
4.4.4.4-
+ + + +
4-4-4-4.
4-4.4.4.
+ + + +
Viseeral (11 cases) -
Late skin (9 cases)
Total
29
Case J-12727. Patient complained of pain over right side of head from
eye to ear, duration four months. A definite area of tenderness to pres-
sure above the zygomatic bone was found. No tenderness over sinuses or
mastoids. Ptosis of right eye, complete inability to move right eye ball.
Pupils equal, circular. Vision of both eyes apparently normal by rough
tests. Fundi, moderate arteriosclerosis. No change in retina of right eye.
No facial paralysis but deviation of the mouth to the right upon showing
teeth. On genitalia or dorsum of prepuce there was a definite indurated
scar 1 centimeter in diameter. Wassermann = Neg. 2. Kahn 4. Diag-
nosis: Syphilis. Periostitis of orbit?
Case G-83908. June 16, 1923, gumma of palate and central nervous
system (VIII nerve) was diagnosed. Under treatment her deafness (right
side) cleared up entirely. Patient received very irregularly three courses
of arsphenamine and two of mercury and potassium iodid. The Wasser-
mann test was positive at all times. February 10, 1928, patient was given
several injections of arsphenamine and bismuth up to January 23, 1929.
January 23, 1929. Wassermann = Neg. Neg. Kahn 4.
During the above second period of treatment the Wassermann became
negative for the first time with ice-box method June 22, 1928, and remained
46,2
Monserrat: Kahn Test in Syphilis
233
so until January 23, 1929, except on the following dates: October 19, 1928,
Wassermann = Neg. 3. January 4, 1929, Wassermann = Neg. 3. Diag-
nosis: Old gummatous perforation of soft palate.
Case 20149. Patient was operated on October 15, 1928, for a myoma of
the uterus (hysterectomy). Very satisfactory post-operative convales-
cence. Came back to the hospital on December 8, 1928. Physical exam-
ination showed an enlargement of heart to the left, apical systoic murmur.
Aortic second sound somewhat coarsened. Murmur not transmitted to ves-
sels of neck. Radial arteries thickened. Blood pressure 115/75 left,
125/85 right. No evidence of syphilitic infection. August 22, 1928, Was-
sermann reaction = Anticomplementary. August 28, 1928, Wassermann
reaction = Positive (water bath = 4; ice box = 2). December 11, 1928,
Wassermann reaction = Negative.
Two successive Wassermann tests done in the laboratory of the hospital,
one on December 11, 1928, and the other on January 10, 1929, using the
ice-box method were also negative. A Kahn test was performed at the
latter date and the result was 4 plus. An X-ray examination showed di-
latation of the aorta and enlargement of the heart.
March 6, 1929, the patient was given an injection of neoarsphenamine
(0.30 gram), and the Wassermann test then became strongly positive
(4-4). The treatment was continued until April 10, 1929. During the
course of this treatment the Wassermann test was constantly 4-4. Im-
pression. Syphilis (seropositive).
Myocardial degeneration with some changes in the aorta (etiology may
be lues, arteriosclerosis; uterine myomata may have some bearing).
Asymptomatic cases, — In this group of asymptomatic cases, as
demonstrated in Table 10, the Kahn test is again more sensitive
than the Wassermann reaction. This shows that the Kahn test
is perhaps a more dependable procedure than the complement
fixation for the diagnosis of chronic syphilitic cases.
Table 10. — Showing the results of the Wassermann and Kahn tests in
thirty-five cases of latent syphilis.
[Very strongly positive, j I f j 1 -f ; strongly positive, +++ ; moderately positive, Hr+ ; slightly
positive +; negative, — .]
Number of cases.
Kahn.
Wassermann.
Water bath.
Ice box.
17 _
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+ + +
+ +
+
+ + + +
+ + +
+ + + +
1_
+ + + +
1
+ + + +
1. _._
+ +
1
— J-2506 (1-23-1929).
2
— H-93343 (1-15-1929).
2 _ _
—
3„_
— H-13182 (XI-7-1928).
7 __
—
234 The Philippine Journal of Science i»ai
Case J-2506. In 1911 at the age of 16, patient had a genital sore
that was thought to be a primary lesion. There was no record of her
having received any antisyphilitic treatment. She came back in 1922
complaining of generalized pruritus. A diagnosis of syphilis (seropositive)
was made at that time. Out of four Wassermann tests performed, two were
positive and two were negative. She received 3 doses of diarsenol and did
not present herself again until August 17, 1928, when she complained of
essentially the same, generalized pruritus. Patient's history was essen-
tially negative, except for shortness of breath on exertion and micturition
(5 or 6 times each night). Blood and spinal Wassermann were negative.
October 12, 1928, patient was given an injection of silver arsphenamine,
0.1 gram. October 19, 1928, the Wassermann showed water bath = nega-
tive; ice box = 1. Another injection October 19, 1928, of silver arsphena-
mine 0.1 gram. December 7, 1929, the Wassermann showed water bath =
negative; ice box = 1. Another injection October 19, 1928, of silver ars-
phenamine 0.1 gram. December 7, 1929, the Wassermann was Neg.-
Neg. The patient was given six injections of arsphenamine (0.20 gram
each), and one injection of bismuth (0.20 gram) up to January 23, 1929.
Repeated Wassermann during this treatment remained negative.
On January 23, 1929, blood was tested again for Wassermann and
Kahn. The results were the following: Wassermann Neg.-Neg. Kahn 4.
Diagnosis: Syphilis Wassermann reaction (1922).
Case H-93343. History of syphilis. February 17, 1928, the Wasser-
mann reaction showed water bath = negative ; ice box = 4. Patient was
given arsphenamine and bismuth and the Wassermann reaction in July,
1928, became completely negative. January 15, 1929, patient came again to
the hospital. This time the Wassermann reaction was found negative, but
the Kahn test positive ( + + + )• At this time, however, there was no
evidence of syphilis on physical examination. Diagnosis: Latent Was-
sermann reaction, late (1928).
Case H-13182. Patient had one premature spontaneous delivery (still-
born baby) on December 4, 1924. Denies venereal infection. Wassermann
= 4-4 on December 16, 1925. Positive. (Patient's husband received an-
tisyphilitic treatment at that time in the dispensary of the hospital.)
Patient came for the second time to the hospital in 1927. Started treat-
ment with silver arsphenamine, neoarsphenamine and bismuth from Feb-
ruary 23, 1927, to January, 1928. During the course of this treatment
the Wassermann remained negative.
November 7, 1928, Wassermann Neg.-Neg, Kahn test 1. Diagnosis:
Latent syphilis (1925).
Syphilis of the central nervous system. — In the twenty-one
cases examined, the Kahn test is undoubtedly more sensitive than
the Wassermann water-bath method.
Congenital syphilis. — In the two cases examined the results
were identical.
46,2
Monserrat: Kahn Test in Syphilis
235
Table 11.—- Showing the results of the Wassermann and Kahn tests in
twenty-one cases of syphilis of the central nervous system and in two
cases of congenital syphilis.
[Very strongly positive,
-+; strongly positive,
positive, + ; negative, — .]
; moderately positive, +4-; slightly
Type.
Number
of
cases.
Kahn.
Wassermann.
Water bath.
Ice box.
Neurorecurrens
1
2
1
1
1
2
3
1
1
1
1
1
3
2
+ + + +
+ + + +
+ + + +
+ + + +
+ + + +
+
+ + + +
+ + + +
+ + + +
+ + +
+ +
+
+ + + +
+ + + +
+ + + +
+ + + +
+
+ + + +
+ + + +
+ + H-811 (XII-10-28).
— H-56170 (1-23-29).
+ + + +
+ + + +
+ + + +
+ + + +
+ +
—H-30276 (XI-5-1928).
+ + + +
General paresis (4 cases) <
Tabes (5 cases)
Asymptomatic and nonspe-
cified (11 cases)
Congenital
Case H-811. Patient in 1924 complained of nervousness, speech trouble,
and weakness. Denied lues and gonorrhea. Had headaches and difficulty
of vision for five years. Pupils irregular. No reaction to light. Marked
tremor of tongue and facial muscles. Marked slowing and slurring of
speech, but test phrases were pronounced quite well when patient really
tried. Tremor of hands. Patient's memory quite good.
Globulin test 4. Wassermann of spinal fluid (positive, 1 cubic centi-
meter). Colloidal mastic, 5555554321.
Patient received treatment with tryparsemide from May 23, 1924,
to January 20, 1925. Patient came back to the hospital December 10, 1928.
At this time Wassermann showed water bath = Neg., Ice box = 2. The
Kahn test = 4. Diagnosis: General paresis.
Case H-56170. Patient was seen first in psychiatric clinic August 17,
1926, with a complaint of loss of consciousness and failing memory.
Globulin 4. Wassermann, water bath = Neg., Ice box = 4. Mastic test,
5543210000.
Treatment began September 15, 1926, with neoarsphenamine-tryparse-
mide and bismuth. Blood Wassermann remained negative as a result of
treatments in October, 1928, up to January 23, 1929.
January 23, 1929, Wassermann Neg. Neg., Kahn 4. Diagnosis: Cen-
tral nervous system syphilis (paresis). Aortic insufficiency.
236
The Philippine Journal of Science
1931
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Monserrat: Kahn Test in Syphilis
237
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238 The Philippine Journal of Science mi
Case H-30276. Patient, seen May 5, 1928, complains of severe head-
aches at menstrual periods and sore throat. No clinical evidence of lues.
One year before patient had miscarriage (in the third month).
May 12, 1928, spinal fluid Wassermann 4-4 (1 cubic centimeter and
0.4 cubic centimeter of fluid). Colloidal mastic test, 2221000000.
Patient started treatment (arsphenamine, neoarsphenamine and bis-
muth) May 1928.
November 5, 1928, Wassermann blood Neg. Neg., Kahn 1.
Influence of treatment upon the Kahn and Wassermann tests. —
The influence of treatment upon the Kahn and Wassermann
tests was studied in 25 cases, which received weekly intrave-
nous injections of arsphenamine and neoarsphenamine. Bis-
muth was also used in cases with secondary and tertiary mani-
festations of the disease. In Table 12, we selected 11 cases
that were fair representations of the different varieties of sy-
philis. The results showed that the Kahn precipitation test
remains positive longer in patients under the influence of various
treatments than either of the complement-fixation reactions.
Therefore, in the presence of a known case of syphilis, a nega-
tive Kahn test will have perhaps a greater diagnostic value than
the Wassermann methods.
SUMMARY AND CONCLUSIONS
Two hundred twenty cases were studied under clinical con-
trol and comparisons were made between the Kahn and the
Wassermann tests. The water-bath and the ice-box methods
with a sensitive antigen were used in the Wassermann reaction,
and the latter method was found the more delicate of the two.
The technic employed in the Kahn test was the latter method,
proposed by its author. The results seem to agree in general
with those obtained by other investigators in regard to the
sensitiveness of the Kahn test. In the few nonsyphilitic cases
examined no false results have been observed, and in the syph-
ilitic cases, the Kahn test has consistently appeared more
pronounced especially in latent syphilis, and in cases under the
influence of treatment, as compared with the Wassermann ice-
box fixation method. Compared with the ordinary water-bath
method, the Kahn test is undoubtedly very much more sensitive.
We feel, therefore, that the Kahn reaction when properly done
and properly interpreted is a valuable test for the serological
diagnosis of syphilis and should be used in routine work in
conjunction with the Wassermann reaction, particularly when
the water-bath method is the only method used. It will give
the physician a more dependable laboratory diagnosis than the
Wassermann water bath alone could give.
46.2 Monserrat: Kahn Test in Syphilis 239
ACKNOWLEDGMENT
I am indebted to Professors Carrol G. Bull and G. Howard
Bailey for their valuable suggestions and for the facilities ex-
tended to me during this work in the laboratory of the School of
Hygiene and Public Health of the Johns Hopkins University,
and to Prof. Allan M. Chesney and Dr. H. H. Hopkins for their
courtesies in permitting the use of material from their clinic
in the Johns Hopkins Hospital.
262412 6
COMPARATIVE SEROLOGIC STUDY OF VERNES,
WASSERMANN, AND KAHN REACTIONS IN
EXPERIMENTAL TREPONEMATOSES
By Carlos Monserrat
Of the Department of Pathology and Bacteriology, College of Medicine
University of the Philippines
In the course of the investigations performed by Dr. Otto
SchobKD and his collaborators on experimental yaws and syph-
ilis in Philippine monkeys, hundreds of these animals were ino-
culated with yaws, syphilis, or both.
Through the courtesy of these investigators, I was permitted
to utilize the blood of some of these experimental animals for
the purpose of studying the sensitiveness of the Vernes method,
as compared with the Wassermann and Kahn methods, and of
determining whether or not the Vernes reaction gives regular
results in yaws- or syphilis-infected Philippine monkeys.
MATERIAL INVESTIGATED
For the information of the reader the following data must
be mentioned:
1. The blood of twenty-one infected Philippine monkeys was
examined and reported for this paper. The majority of these
animals had been inoculated within the last five years.
2. The strains of Treponema pertenue were secured by direct
inoculation from patients in the Philippines to monkeys, and
were maintained alive through successive passages in monkeys.
3. The strain of Treponema pallidum used in these monkeys
was the well-known laboratory strain known as "Nichols
strain."
4. Some of these monkeys had received some intramuscular
injections of neosalvarsan in the past, and a few had received
injections of heated antitreponematous vaccines.
5. In this investigation ten normal Philippine monkeys were
tested as controls. The blood of these normal animals invariably
gave negative results with the Vernes, Wassermann, and Kahn
tests.
241
242 The Philippine Journal of Science 1931
TECHNIC
During this investigation the Wassermann and Kahn tests
were performed by Dr. Onofre Garcia, of the biologic division,
Bureau of Science, on the day following the bleeding of the
animals. The technic for the Wassermann test was the same
as that described previously by Dr. Otto Schobl and the writer ;
(2) that is, guinea pig's complement, antimonkey hemolytic
system, and cholesterinized antigen.
The technic followed in performing the precipitation test is
the standard method of Kahn. (3)
The samples of blood for the Vernes test were received by
the present author at irregular intervals, and the Vernes reac-
tion was performed with the sera at periods of from two to ten
days after the bleeding of the animals.
The Vernes reaction was performed by following exactly
Professor Vernes's technic, which the author of this paper
learned in the laboratory of Professor Vernes, at the Prophy-
lactic Institute of Paris, during his last trip to France in the
autumn of 1929.
The results of the Wassermann and Kahn tests are as follows :
Very strongly positive -f + + + (100 per cent haemolysis).
Strongly positive -| — j — 1_ (75 per cent haemolysis).
Moderately positive -f--f- (25 per cent haemolysis).
Slightly positive + (10 per cent haemolysis).
Very slightly positive (doubtful) ± (5 per cent haemolysis).
Negative — (no haemolysis).
The results of the Vernes reaction in our tables are given in
figures that exactly represent the numbers of the photometric
readings of each sample of blood. In this way, and following
the advice of Professor Vernes, misinterpretations were avoided
in the results of the Vernes reaction when compared with the no-
menclature generally adopted in the readings of the Wassermann
and Kahn tests.
Since the Vernes method gives to the clinician a more quan-
titative measurement of the treponematous infection in the
patient than either the Wassermann or the Kahn test, because
the reading in the first test is made by means of a photometer,
the table adopted by Vernes for the clinical interpretation of
these figures is also given here.
The following table is based on a great number of clinical and
serologic studies of normal and syphilitic patients.
46,2 Monserrat: Serologic Studies 243
SYPHILIMETRIC TABLE
Photometric reading -0. This means a completely normal serum.
Photometric reading 1-2. This means a normal serum, although some-
what doubtful.
Photometric reading 3-4. In 100 normal sera, there were only 2 sera
that gave the values 3-4. In 100 sera taken at random approximately 25
syphilitic sera were found which gave the values 3 and 4. The formula is
25-3
75 X N
Photometric reading 5-6. In 500 normal sera, only one serum gave the
values 5-6. In 100 sera taken at random approximately 50 syphilitic and
5Q 3
50 non-syphilitic sera gave the values 5 and 6. The formula is — •
50 X N
Photometric reading 7, 8, 9, 10, 11. In 2,000 sera there were approxi-
mately 1.999 syphilitic sera and only one non- syphilitic serum. The for-
1.999 - S
mula is
1-N
Photometric reading 12, 13, 14, 15, 16, 17, 18. In 10,000 sera there
were 9.999 syphilitic sera and only one non-syphilitic serum. The for-
, . 9.999 -S
mula is— - =r^-.
1 — N
Photometric reading 19, 20, 21, 22, 23, 24, 25, 26, 27. In 650,000 sera
there were approximately 649.999 syphilitic sera and only one non-syphili-
649.999 S
tic serum. The formula is ' - — .
1 — N
Values higher than the photometric reading 27. Indicate syphilitic in-
fection without exception. The formula is S — sure.
Taking into consideration the results obtained with the
Wassermann reaction, our twenty-one sera examined here were
classified as follows :
Number
of sera.
Slightly positive + 7
Moderately positive ( + + ) 2
Strongly positive ( + + + ) and ( + + + + ) 12
Total 21
RESULTS OF TESTS
The results of the tests presented in Table 1 show that the
Vernes method follows the results of the Wassermann test more
closely than those of the Kahn test, with the exception of monkey
12, and that in the slight reactions the Vernes test gives more
definite and clear-cut results than the Wassermann test itself.
In regard to the Kahn test, Vernes reaction is also more sen-
sitive in the slight reactions.
244
The Philippine Journal of Science
1931
Table 1. — Showing the results of the blood in seven infected Philippine
monkeys.
Monkey.
Test.
Num-
ber.
6
7
8
9
10
11
12
Designation.
Wasser-
mann.
Yae-10
Sy-D-20
F-38
Ym-20 no clip
L-13right
Sy-3 ___
K-28
4-
4-
4-
4-
4-
4-
4-
Kahn.
4-4-
4-
±
4-
Date
performed.
7~ 1-30
7- 7-30
7- 7-30
7-23-30
7-23-30
6-24-30
7- 9-30
Vernes.
4
10
11
6
5
10
0
Date
performed.
7-11-30
7-11-30
7-11-30
7-29-30
7-29-30
6-26-30
7-13-30
Table 2 shows the results of the tests of two moderately posi-
tive sera. The Vernes reaction here also seems to be more
sensitive than the Wassermann, more especially in monkey 13,
and more sensitive than the Kahn test in monkey 14.
Table 2. — Showing the results of the tests in two moderately positive
sera.
Monkey.
Test.
Num-
ber.
Designation.
Wasser-
mann.
Kahn.
Date
performed.
Vernes.
Date
performed.
13
14
L— 15 cut tail
4-4-
4-4-
4-44-
-f-
7-23-30
11- 4-30
22
7
7-29-30
9-11-30
W-23__
Table 3. — Showing the results of the Vernes, Wassermann, and Kahn tests
in strongly positive sera from infected Philippine monkeys.
Monkey.
Test.
Num-
ber.
Designation.
Wasser-
mann.
Kahn.
Date
performed.
Vernes.
Date
performed.
15
16
17
18
19
20
21
22
23
24
25
25
Yac-12
4-H-4-4-
4444
4-4-4-4-
4-4-4-4-
4-4-4-4-
4- + 4-4-
4-4-4-4-
4-4-4-4-
4-4-4-4-
4-4-4-4-
4-44-4-
4-4-4-
4-4-4-4-
4-4-4-4-
4-4-4-4-
4-
4-4-4-4-
+ + + +
4-4-
4-4-4-4-
4-4-
4-4-
7- 1-30
7- 1-30
7- 7-30
7-24-30
7-24-30
7-24-30
6-20-30
8- 6-30
8- 6-30
8- 8-30
10-30-30
11- 4-30
94
60
62
32
49
76
91
67
89
136
25
14
7-11-30
7-11-30
7-11-30
7-29-30
7-29-30
7-29-30
6-26-30
8-13-30
8-13-30
8-13-30
11- 4-30
11-11-30
B-9 J
E-14-instr. tail—
O-C both clip
Sy_p_23
K-13-left
Yaw-V-10
F-2 _
J-l __
G-25
J-ll
W-25
46'2 Monserrat: Serologic Studies 245
Table 3 shows the results with strongly positive sera from
twelve infected Philippine monkeys. The Vernes reaction fol-
lows again more closely the results of the Wassermann rather
than the Kahn test and gives a better measure of the amount
of the treponematous "reagin" in vivo.
SUMMARY
The blood of twenty-one Philippine monkeys infected at dif-
ferent intervals of time with yaws, syphilis, or both, have been
tested. A few of these animals received neosalvarsan treat-
ments in the past and also injections of heated antitrepone-
matous vaccines.
The results of the Vernes, Wassermann, and Kahn tests agree
in a general way, but the Vernes reaction follows more closely
the results of the Wassermann test in spite of the fact that the
Vernes reaction was performed a long time after the bleeding of
the animals. This circumstance necessarily will bear some in-
fluence on the results and the sensitiveness of the test. Never-
theless, in our series the Vernes test is found somewhat more
sensitive than our Wassermann test.
Since the readings of the Vernes test are made by means of
a photometer and the results are expressed in figures, a more
accurate quantitative measurement of the "reagin" is made
possible. The results, therefore, are more helpful especially for
the clinical interpretation of weakly positive and border line
results for which the reading of the Wassermann and Kahn
tests are usually insufficient.
It must be borne in mind that the precipitation reactions in
Philippine monkeys give much lower results than the Wasser-
mann test, as proven by Doctors Schobl and Garcia with the
Kahn test. Nevertheless the Vernes reaction, being a precipita-
tion reaction, gives higher values, or did in our experiments,
than both the Wassermann and Kahn tests.
CONCLUSIONS
1. In Philippine monkeys inoculated with yaws or syphilis,
the Vernes reaction was found regularly positive.
2. This fact shows the sensitiveness of the reaction of Vernes,
which is a precipitation reaction. It is known that precipita-
tion reactions (Kahn) are not as pronounced in Philippine
yaws or syphilitic monkeys as the Wassermann test.
246 The Philippine Journal of Science
3. This is particularly evident in sera with a low and a mod-
erate degree of positive reaction (Table 1), whether compared
with Wassermann or Kahn reaction.
4. Sera from infected Philippine monkeys giving high positive
values with Wassermann reaction likewise give high values
with Vernes reaction, unlike those with Kahn.
5. With regard to Philippine monkeys the Vernes reaction
seems to have an advantage over both the Wassermann and Kahn
tests.
REFERENCES
1. Schobl, Otto. Philip. Journ. Sci. 35 (1928) 209.
2. Schobl, Otto, and Carlos Monsbrrat. Philip. Journ. Sci. § B 12
(1917) 249.
3. Kahn, R. L. Serum diagnosis of Syphilis by Precipitation; governing
principles, procedure, and clinical application of the Kahn precipi-
tation test. Williams and Wilkins Co. (1925).
MALARIA TRANSMISSION IN THE PHILIPPINES, IV
METEOROLOGICAL FACTORS1
By C. Manalang
Of the Philippine Health Service, Manila,
THREE TEXT FIGURES
Mayne, 2 in his article on the influence of relative humidity on
the presence of malarial parasites in the insect carrier, mentions
the work of Bentley on the influence of temperature and humid-
ity on the malaria incidence of Bombay between 1909 and 1911,
which brought to light a definite relationship between the months
of heaviest infections and the phenomenon of relative humidity.
Bentley found that the occurrence of new infections coincided
with a period of slightly lower but not more uniform high tem-
perature in the presence of increased humidity. Mayne' s work
covered dissection of 5,052 mosquitoes from March to Septem-
ber, 1927, in the District of Saharanpur, United Provinces, India.
Out of this number, he found five infected A. calicifacies — Giles
(total number of this species, 2021) collected from August 9
till September 8, a period with the highest relative humidity
(82 to 99 per cent). Gill,3 in his epidemiological methods of
forecasting seasonal appearance of endemic or epidemic malaria
in Punjab, India, uses biological as well as meteorological fac-
tors. Wenyon4 believes that in the natural infection of mos-
quitoes, temperature is a much more important factor than
humidity. He asserts that there is no evidence that the effects
of humidity of the atmosphere play any part in the active devel-
opment of parasites to the mosquitoes. "Provided there is suf-
ficient moisture in the air to enable the mosquito to live, the
malarial parasites will develop normally.,, He agrees with
1 From the field laboratory, division of malaria control, Philippine Health
Service, Tungkong Manga, Bulacan. The writer expresses here his grati-
tude to Father Miguel Selga, director of the Weather Bureau, for his
personal interest, valuable suggestions, loan and installation of instruments,
and training of the laboratory personnel in making observations.
2 Indian Journ. Med. Res. 15 (1928) 1073.
8 Cited from Mayne.
4 Cited from Mayne.
247
248 The Philippine Journal of Science 1931
Gill that the spread of malaria may, however, be affected by
lack of humidity, but only on biological grounds, because the
mosquitoes which ingest parasites may not live long enough for
sporozoites to appear in the salivary glands. In Europe (North
Holland)5 malaria in mosquitoes is prevalent in autumn and
winter with its maximum in November or December, while ma-
laria in man is a phenomenon of spring and summer with its
maximum in June or July. Swellengrebel (p. 25) says:
The fact that in two localities so near each other as Nieuwendam and
Wormerveer or Sloten, there is no synchronism in the epidemic period-
icity (beginning of epidemic in Nieuwendam in 1912 with remission in
1914-1917, at Wormerveer beginning of epidemic in 1918, at Sloten in
1921) makes it doubtful whether climatic conditions can have much in-
fluence. On the other hand, the synchronous decline of the epidemic in 1923
indicated the presence of a common inhibiting factor. Was this factor
the low temperature of the fourth quarter of 1922 and the first and sec-
ond quarters of 1923? If so, why did the epidemic cease at Wormerveer
after 1902 and why did it show a remission in Nieuwendam? Have the
dryness and high temperature anything to do with it; if so, why did not the
climate in 1911 produce a similar effect? (The following year witnessed an
exacerbation of epidemic at Nieuwendam.)
These considerations make it impossible to attribute any epidemiolog-
ical importance in the climatic changes observed here, the more so as no
influence can be detected on the anopheline population.
Granting that the epidemics were diagnosed correctly, Swel-
lengrebel's conclusions are to be expected but his data are sub-
ject to further analysis, for, (1) to expect synchronism in the
epidemics between 1912 and 1921 in the three nearby places due
to a common cause (climate) several variable factors, as the mos-
quito density, the number of suitable human carriers, and their
accessibility to the mosquito, including those of the susceptibles,
should be equally present in all. (2) The introduction of a new
parasite strain in one place and not in the others should be con-
sidered. On the other hand, the synchronous decline of malaria
in 1923 cannot be explained by such variable factors as coincident
decrease in the number of suitable carriers, immunity, treatment
of cases, mosquito control, improved living conditions, etc., all
happening at the same time in the three places, all the more so
when the campaign in these places from 1920 to 1923 was limited
to mosquito control measures, and at Nieuwendam, this work was
confined to catching adults in the stables only (pp. 31-35) . Since
• Principles and Methods of Antimalarial Measures in Euro^:2d
general report of the Malaria Commission, League of Nations (19,27) bl.
Malaria in the Kingdom of the Netherlands (1927) 67-70, graphs 4
and 5.
46,2 Manalang: Malaria Transmission 249
climatic records are available from 1902 to 1923, while the ano-
pheles data are only available for the years 1920 to 1923, the role
of the transmitter under the existing climatic conditions in the
incidence of malaria from 1902 to 1919, inclusive, is not known.
The trend of the disease during this period cannot and should not
be explained on the basis of the mosquito findings from 1920 to
1923 alone. It would seem, therefore, that in the synchronous
decline of the epidemic in 1923, with a parallel trend of the
disease from 1921 to 1923 in the three places mentioned, (graph
5, p. 70) while no climatic influence was detected on the anophe-
line population, one cannot eliminate entirely a common inhib-
iting factor (the low temperatures in the fourth quarter of 1922
and the first and second quarters of 1923). Neither can the
influence of the climate on the behaviors of the disease and on the
mosquitoes previous to 1920 be ignored because anopheles and
human carrier data are not available or have not been utilized.
One point is indisputable in SwellengrebeFs observations, and
that is the coincidence of malaria in mosquitoes with the months
of high relative humidity (see his table 4a, p. 45 and graph 6,
P. 71).
The results of two years (September, 1927, to August, 1929)
systematic captures and dissections for natural malaria infection
of A. funestus Giles in two adjacent camps of La Mesa and
South Portal of the Novaliches water project,6 form the basis
of the present article. Meteorological records were taken from
the field laboratory at Tungkong Manga, 9 to 10 kilometers north
of La Mesa and South Portal but at about the same elevation
(100 meters) above sea level. Unfortunately, these observations
were not carried on simultaneously with the mosquito observa-
tions, but were from September, 1929, to August, 1930, and may
differ from those obtaining from September, 1927, to August,
1929. The average monthly rain gauge readings from Septem-
ber, 1927, to August, 1929, at La Mesa, however, show the same
proportionate distribution as those from September, 1929, to
August, 1930, at Tungkong Manga, although somewhat lower.
Any difference in temperature and relative humidity between the
two periods at the two places would probably be a difference in
degree only and not in distribution. The use of meteorological
data for September, 1929, to August, 1930, therefore, seems
justified and should give at least a relative value.
"See the preceding three papers.
250
The Philippine Journal of Science
1931
800
700
600
£500
"cJ400
= 300
^200
100
0
3
2 c
O
1 t
o°-
A
Ra
nfc
ill-
/\
/
' \
N
/
/
\y
*/ N.J
^
/
\\—
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s~
s —~
V
/ V
""""*\
' —
.—.''/~
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iva
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y*
Fig. 1. Showing a rise of the rates of infection in Anopheles funestua during the period
of heaviest rain although the infections were also present during the dry months.
28
27
o>26
|25
J?24
§23.3
(0
Of
o
1
0
-Ten
iper
atu
»e
-*\
\
\
\
7
~~/
Sal
var
fgh
»nd-
p
-5*
omc
ch
*~*
7\
[/ N.
^7
t
— "v
>
- -r'"
•— ■
V
^W
^
.-•'*
s%
fe o *o ajo § -= p a) u o a)
Fig. 2. Showing a rise of the rates of infection in Awopfeefes funestus during the warmest
months although they were also present during the cool months.
46,2
Manalang: Malaria Transmission
251
88
85
8E
79
"£76
°73
c
a»
z
1
0
Rel
ativ
a hi
mid
iiy-
\
,
/
/
'
. /
Sal
vap
^l<
and-
/
/
\
-s
tarn
ach
-'
-"
/
/ \
/
V
' »». *
\
\
•
—
y
V
"^.^
•
-j u- ^ <l s — j — ) < (o o Z a
Fig. 3. Showing some rise in the rates of infection in Anopheles funestus during the months
of high relative humidity although they were also present in March and April, months
with the lowest percentage of relative humidity.
Table! 1. — Dissection of funestus from La Mesa and South Portal.
Month.
January...
February..
March
April
May
June
July
August
September.
October
November.
December.
Number
dissected.
932
327
878
668
686
782
883
571
1,058
1,190
767
803
Positive
stomachs.
11
6
15
4
18
8
23
17
8
12
8
14
Per cent.
1.2
1.8
1.7
0.6
2.6
1.0
2.6
3.0
0.7
1.0
1.0
1.7
Positive
salivary
glands.
Per cent.
0.8
0.9
1.6
1.3
2.8
1.4
1.1
1.2
0.7
0.1
0.5
1.1
Table 1 shows the monthly catches and infections found during
the period of observation. The influence of larval control on the
adult density and uncontrolled movements and quininization of
252
The Philippine Journal of Science
1931
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46,2 Manalang: Malaria Transmission 253
the population on mosquito infection are not known. Tempera-
ture shows no apparent influence on breeding and adult density
in this locality.
Table 2 shows the monthly mean meteorological observations.
COMMENTS
Given a favorable adult funestus density, a community of the
topography of South Portal with a low rate of suitable human
carriers would be expected to have a seasonal prevalence of ma-
laria coinciding with the months of most rainfall, high mean tem-
perature and relative humidity. Areas where the funestus
breeding streams dry out during the dry months, would be ex-
pected to have malaria during the rainy season. On the other
hand, localities where the breeding is limited to permanent
streams, heavy rains would flush the larvae, reduce the density
and transmission even in the presence of suitable carriers.
Since funestus has been found infected in nature in all months
of the year provided suitable carriers are available, the observed
prevalence of malaria during the rainy season in one region and
dry season in another, may be explained by the influence of the
rain on the amount of breeding and the resultant adult density
of the transmitter. Both types of breeding, permanent and tem-
porary streams, exist in the Novaliches water project, as pre-
viously mentioned,7 and explains the uniform high funestus den-
sity observed in the camps in 1927 and 1928. The natural
decline in funestus density observed in South Portal in 1929
and 1930, and in North Portal and Tungkong Manga in 1929,
(the former dropping in February, the latter in May) cannot
be explained.8
SUMMARY
1. From the available mosquito data at the La Mesa and South
Portal camps of the Novaliches water and meteorological data
at the field laboratory in Tungkong Manga, malarial infection
in A. funestus Giles shows a rise in the rates with the increase
of rainfall, mean temperature and relative humidity, although
infections were also present during the dry, cool, and less humid
months.
2. Anopheles funestus breeds in permanent or temporary
streams and the influence of the rainy season on breeding and
the resultant adult density probably explains the different sea-
sonal distribution of malaria transmission in the Philippines.
7 Manalang, Philip. Journ. Sci. 37 (1928) 123.
8 See preceding article.
ILLUSTRATIONS
TEXT FIGURES
Fig. 1. Graph showing a rise of the rates of infection in Anopheles funes-
tus during the period of heaviest rain although the infections
were also present during the dry months.
2. Graph showing a rise of the rates of infection in Anopheles funes^
tus during the warmest months although they were also present
during the cool months.
3. Graph showing some rise in the rates of infection in Anopheles fu-
nestus during the months of high relative humidity although they
were also present in March and April, months with the lowest
percentage of relative humidity.
2S2412 — 7 255
LEAF AND SEED STRUCTURE OF A PHILIPPINE
CORIARIA
By Jose K. Santos
Of the Department of Botany, University of the Philippines
and of the Bureau of Science, Manila
FOUR PLATES
This rare shrub, a species of Coriaria, occurs in the Mountain
Province, northern Luzon, and is the only species recorded in
the Philippines under the small family Coriariacese. It is bota-
nically known as Coriaria intermedia Matsumura. For a num-
ber of years this plant has been a subject for research on account
of the reputed poisonous properties of its leaves and seeds.
Chemical and toxological investigations have been made and at
present a still more extensive research on the same line is being
conducted. Very interesting results have already been obtained.
This critical study of the structure of the leaf and seed of the
Philippine Coriaria was undertaken, therefore, with a view to
having a definite basis for the identification of fragments of the
plant in cases of poisoning. Then, too, it was thought that this
plant, being rare in this country, might show some structure
useful to the systematic anatomy of the Phanerogams.
The late Eduardo Lete, pharmacist from San Fernando, La
Union, was the first to report the poisonous properties of Phil-
ippine Coriaria to the Bureau of Science about the year 1915.
During the summer of 1916, while the writer was detailed in
Benguet by the Bureau of Science to collect some botanical spe-
cimens, he obtained direct information from some of the natives
in Trinidad valley, Baguio, concerning the poisonous effects on
man of this plant, commonly known by them as buakat, or
backet. He was told! that at one time a certain couple with
their two children died o^ poisoning after taking a decoction of
the fruits and leaves of bvjakat or backet, which they mistook
for their native Benguet tea, because of the similarity of the
two plants. In the summer of 1918, a year later, the writer
was assigned to conduct a more extensive botanical exploration
in Haight's Place and vicinity, about 60 miles north of Baguio.
He collected several kilos of the fruits and shoots of Philippine
257
258 The Philippine Journal of Science i«>3i
Coriaria and from this material attempts to isolate its active
principle and to conduct experiments showing its action upon
animals were made by the chemists of the Bureau of Science.
In 1919, Wells (7) reported that Coriaria intermedia contains a
poisonous glucoside in its leaves and fruits.
Lindsay (2) claims that in New Zealand there are at least
three species of Coriaria; among them is Coriaria ruscifolia Lin-
naeus, the most abundant and popularly known as toot-poison be-
cause of its poisonous properties. The action of the poisonous
portions of this plant on man, cattle, and on sheep were de-
scribed by him respectively. He indicated that toot-poison
belongs to the class narcotico irritants.
Among the known species of Coriaria, the only one that has
been thoroughly investigated is Coriaria myrtifolia Linnaeus.
This species is found distributed in the southern part of France,
Spain, and Italy. According to Reutter(5) it contains a gluco-
side called coriamyrtin and an alkaloid coriarine, also a con-
siderable amount of tannin and resinous substances. Its im-
portant anatomical features described by Solereder(6) may be
summarized as follows : (a) The stomata occur on both surfaces
of the leaf and they are adjoined on either side by a single sub-
sidiary cell, parallel to the pore; (6) the upper and lower epi-
dermal cells in surface view are polygonal in outline; (c) the
leaf tissue is nearly centric and almost entirely formed by pali-
sade tissue; (d) the outer limit of the bast is formed by massive
isolated groups of bast fibers; (e) the medullary rays are broad
and are as much as seven cells in breadth, and the medullary cells
are elongated in a vertical direction; and (/) the end-walls of
the vessels have simple perforations and the wood parenchyma
has simple pits.
Recently Kariyone and Sato(l) reported that Coriaria japo-
nica A. Gray, contains coriamyrtin, similar to the one isolated
from Coriaria myrtifolia.
MATERIAL AND METHODS
The seeds and leaves used for this study were collected by the
author last summer from plants growing in the city of Baguio
and along the trail leading to Mount Santo Tomas. The fruits
and leaves were preserved in 6 per cent solution of formalin.
The study of the flowers was made from the dry specimen kept
in the Bureau of Science and from the fresh material that was
generously sent to the writer by Mr. Sixto Laraya, District
Forester, stationed at Baguio.
46»2 Santos: Philippine Coriaria 259
The seeds are very minute and enclosed by a very hard peri-
carp, which serves as seed coat and makes sectioning quite
difficult. This difficulty was remedied, however, by embedding
them in a thick paste of gum arabic, which was subsequently
exposed at room temperature until the consistency of the gum
was suitable for sectioning. Several free-hand sections were
made by using a gillette blade. The cross section through the
blade was prepared by means of a sliding microtome, stained
with safranin and contrasted with Delafield's hematoxylin and
mounted in balsam.
DESCRIPTION
The plant — This shrub was first described by J. Matsumura
(3) from the specimen collected from Formosa as follows:
Coriaria intermedia, Matsumura, sp. nov. Frutex polygamo-monoicus,
foliorum forma et magnitudine C. Japonicae, A. Gray, similis, antheris
verruculosis, carpellis, versus latus reticulatis inter C. myrtifoliam, L. et
C. nepalensem, Wall, mediatus. Racemi quam eas C. Japonicae, A. Gray,
breviores, 50-90 mm. longi, aphylli, vel foliati. Sepala ovalia, margine
purpures suffusa. Fl. steril. petala minutissima; stamina 10; antheris
oblongo-ellipticis, verruculosis; vestigio germino mullo. Fl. fern; petala se-
palis multo breviora, oblonga, acuta, intus carinata, stamina 10; carpella
5, petalis paulo breviora, matura vix 4 mm. longa, dosali unicostata, versus
latus ventrale prominente venosa.
As to the identity of the Philippine Coriaria, Merrill (4)
reported the following:
Luzon, Province of Benguet, Suyoc to Paui, (4800 Merrill), Nov. 7,
1905. In ravines at about 2,000 m. Formosa.
Specimens of the above number were sent to Dr. J. Matsumura of the
Botanical Institute, Imperial University, Tokyo, Japan, for comparison
with the type of his Formosan species, and after comparing the speci-
mens, he expresses the opinion that the Luzon plant is identical with his
Coriaria intermedia. Specimens collected in Benguet by Vidal, and re-
corded by him as "C. sp. (aff. C. Japonica A. Gray) " are undoubtedly re-
ferable to Coriaria intermedia Matsum. The thirteen known species of the
genus have a peculiar geographical distribution extending from the Me-
diterranean region to the mountains of British India, Japan and Formosa
and from New Guinea to New Zealand, Mexico and South America. The
presence of this Formosan species in Benguet adds another very charac-
teristic species to the known northern element in the Philippine flora.
The Coriaria from the Philippines as observed by the writer
has the following features. It is a shrub from 1 to 3 meters
high. The young branches are quadrangular and of a reddish
or pinkish color with slightly elevated boatshaped lenticels.
The leaves are from 2 to 4.2 centimeters in width and from
4 to 8.5 centimeters in length (Plate 1, fig. 1). They are sim-
260 The Philippine Journal of Science 1931
pie, ovate-lanceolate, trinerved, glabrous and short petiolate
with entire margin. The upper surface is dark green and the
lower surface is light green or sometimes yellowish-green. The
base varies from obtuse to rounded and the apex is acute or
sometimes acuminate. The petiole is very short, from 1 to 2
millimeters long, nearly cylindrical with a shallow groove in
the upper part. The midrib is prominently projecting on the
lower side, at the base of which or directly from the upper end
of the petiole two primary veins arise, one on each side, extended
toward and close along the margin of the leaf and converging
toward the apex. The flowers are arranged in simple racemes,
from 6 to 15 centimeters long and they are provided with bracts
(Plate 1, figs. 2, 3, and Plate 4, fig. S3 a-c). They are poly-
gomo-monoecious, very minute, measuring about two millimeters
in length, and are greenish in color tending to reddish or pur-
plish coloring at the margin of the sepals, or appearing en-
tirely red. The calyx consists of five persistent ovate sepals,
concave in the inner part with acute or acuminate apex, two of
which are slightly smaller in size ; the petals are also five ovate,
very minute, persistent, cream- white in color, with a prominent
angular projection on the inner side (Plate 1, figs. 5, 8) . During
the maturation of the fruit, these petals develop unusually fast.
They cover the cocci and become fleshy and are of a reddish color
turning finally to bluish black, as represented on Plate 1, figures
10 and 14. The androecium of the sterile flower consists of 10
stamens, and that of the bisexual flower varies from 5 to 10.
The anthers of the fully developed male flowers are oblong, ver-
rucous, quadrilocular, introrse and purplish or reddish in color
with long filaments (Plate 1, figs. 7, 9). They measure about
2.2 millimeters in length. The anthers of the bisexual flower
vary from ovate to oblong-ovate or oblong, and usually are
much smaller and have shorter filaments than those of the nor-
mal male flower. They measure from 0.7 millimeter to 1 milli-
meter in length (Plate 1, figs. 4 to 6, 8, 10, and 15) . The gyn-
secium is composed of five more or less independent pistils, with
filamentous reddish or purplish stigmas covered with papillose
appendages (Plate 1, figs. 5, 8, 10). The fruit is composed of
five very small crustaceous cocci, surrounded by fleshy persist-
ent petals and sepals of a bluish-black color, which makes it
berrylike in appearance (Plate 1, figs. 10, 13, and 14).
Structure of the leaf. — The transverse section of the leaf of
Philippine Coriaria is bifacial. The blade is nearly uniform,
46'2 Santos: Philippine Coriaria 261
measuring about 0.25 millimeter. The upper epidermis is com-
posed of a single layer of flattened or rectangular cells with very
thick and highly cutinized outer cell walls. The lower epider-
mis also consists of a layer of cells of the same shape as the up-
per epidermal cells, but they are slightly thinner, their outer
walls are less cutinized and some of them are modified into
guard cells. The mesophyll is differentiated into palisade and
spongy regions. The palisade chlorenchyma occupies about one-
third of the mesophyll and consists of two layers of tubular cells
arranged perpendicularly with distinct intercellular spaces. The
palisade cells of the upper layer are longer than those of the low-
er one. They measure about 0.05 millimeter in length, whereas
the lower palisade cells measure only about 0.03 millimeter.
The spongy chlorenchyma region is made of parenchyma cells
of various forms and sizes, but most are slightly elongated in
the direction parallel to the surface of the leaf. It is richly
supplied with air spaces. Plate 3, fig. 30, represents a trans-
verse section through the midrib showing the character of the
mesophyll described above. Calcium oxalate crystals and epi-
dermal outgrowth are wanting.
In the surface section, the upper epidermal cells are poly-
gonal in outline with from 5 to 7 straight thick walls. They
are from 0.02 to 0.04 millimeter in length and from 0.015 to
0.03 millimeter in width and they are characterized by a fine,
wavy striation that runs either parallel to the longer side of
the epidermal walls or obliquely to the longest side of the epi-
dermal wall. Plate 3, figure 31, is a small portion of the sec-
tion prepared from the upper epidermis showing the surface
view of the epidermal cells with the characteristic striations
of their cuticle. Unlike the upper epidermis of Coriaria myr-
tifolia the stomata are wanting in the upper epidermis. The
surface view of the lower epidermal cells is represented on the
same plate, figure 32. The lower epidermal cells are also poly-
gonal in outline measuring from 0.015 to 0.045 millimeter in
length and from 0.01 to 0.03 millimeter in width, and are char-
acterized by fine striations but their walls are thinner and vary
from four to seven in number. The stomata are somewhat char-
acteristic and numerous. They are not uniform in size and vary
from 0.025 to 0.028 millimeter in length and from 0.012 to 0.015
millimeter in width. They are usually surrounded by four
neighboring cells. Two of these neighboring cells limit the up-
per and lower ends of the stomata while the other two limit
262 The Philippine Journal of Science 1931
the lateral sides and are applied parallel to the length of the
guard cells. The first two are larger in size than the others.
The midrib is convex above and strongly convex below. The
upper epidermis as well as the lower one consists of a single
layer of cells, rectangular, or barrel-shaped, or nearly square in
outline (Plate 3, fig. 30). The outer cell walls of the upper
epidermis of the midrib as well as those of the upper epidermis
of the blade are comparatively thicker than those of the lower
epidermis. The collenchyma cells are poorly developed and as
usual are found in two regions, one just in the inner side of the
upper epidermis above the meristele and the other located within
the lower epidermis below the meristele. The chlorenchyma
cells in the upper region, as well as those of the lower region,
consist of 3 to 4 layers of cells with more or less uniformly and
slightly thickened walls. The cortical parenchyma located be-
tween the meristele and the lower collenchyma is composed of
4 to 6 layers of large polygonal isodiametric thin-walled cells,
while the cortical parenchyma found in the inner part of the
upper collenchyma region consists of three to seven layers of
small polygonal thin-walled parenchyma cells.
The endodermis is somewhat conspicuous. It consists of a
single layer of rectangular, square or polygonal, thin-walled
parenchyma cells. Within this endodermis, the meristele is
located. It is more or less lenticular in shape and the conduct-
ing tissue is somewhat plano-convex in outline. The upper
part as well as the lower part is bounded by two groups of poorly
developed sclerenchymatous cells. The walls of these cells are
not much thickened nor highly lignified. The xylem region is
limited on both the upper and lower part (nearly surrounded)
by small and not distinctly differentiated phloem cells. It is
made up mostly of xylem vessels, from 0.01 to 0.02 millimeter
in diameter and wood parenchyma.
The seed. — The seed is campylotropous, exalbuminous and
inclosed by a hard pericarp. It is kidney-shaped, laterally com-
pressed, more rounded on one margin and the apex narrowly
rounded (Plate 1, fig. 17 a-c) . It measures from 2.2 to 3.2 milli-
meters in length, 1.6 to 2 millimeters in breadth and 1.2 to 1.5
millimeters in thickness. Externally it is brown in color and its
surface is characterized by one prominent dorsal angular eleva-
tion or riblike structure that extends from the upper end of the
hilum or micropylar to its lower or chalazal end, and two to
four elevations or ribs on each lateral or flattened side. These
ribs run parallel to the dorsal one and they are more or less
46,2 Santos: Philippine Coriaria 263
concentric to the hilum. They sometimes anastomose each other
by a few transverse elevations that connect one with the other.
The hilum is somewhat arrow-shaped with the narrow end
toward the apex. The pericarp is hard, cutinized, and lignified.
It takes the place of the outer seed coat or testa. The seed coat
proper is only one and is very thin. The embryo is slightly
bent and whitish in color. The hypocotil is short, conical, and
measures about 0.4 to 0.6 millimeter in length. The cotyledons
are fleshy, plano-convex, and sometimes they are slightly un-
equal in size. The plumule is inconspicuous.
Microscopical structure. — A diagrammatic representation of
the transverse section of the coccus of Coriaria intermedia
drawn under the camera lucida is indicated in Plate 1, fig. 18,
showing the pericarp, the seed coat and a pair of cotyledons.
The section is ovate in outline with a prominent angular protu-
berance at the upper part corresponding to the principal rib
extended along the dorsal side of the coccus, it is more or less
convoluted on the ventral side, corresponding to the section of
the hilum, and is irregularly crenate or wavy on the lateral sides.
The pericarp is differentiated distinctly into two regions, namely,
the parenchymatous region and the stony region. The paren-
chymatous region corresponds to the exocarp and is made up
of several layers of irregularly shaped parenchyma cells with
slightly suberized cell walls which usually contain a brownish
substance. In the outer part it is limited by a single layer of
thick-walled cutinized and somewhat rectangular epidermal cells.
The stony region is differentiated into two parts. The outer
part, which corresponds to the mesocarp, is built up of obliquely
or tangentially arranged elongated stone cells with very thick,
lignified, and pitted cell walls. This portion is more definite on
the lateral sides in which tangentially elongated sclerenchy-
matous elements conspicuously run parallel to the inner surface
of the pericarp. The sclerenchymatous cells of the same re-
gion toward the dorsal sides, however, are mostly arranged
obliquely, following the outline of the outer surface of the peri-
carp. At the region where the elevations or ribs, indicated
above, are located groups of greatly elongated sclerenchy-
matous cells are observed. These sclerenchymatous cells in
cross section appear polygonal in outline with from five to eight
thick, lignified, and pitted cell walls with very much reduced cell
cavity. They measure from 0.3 to 0.4 millimeter in length. The
inner portion of this stony region, corresponding to the endocarp
of a fleshy fruit, consists of radially elongated polygonal stone
264 The Philippine Journal of Science im
cells with greatly thickened and highly lignified cell walls and
with very much reduced cavities. Their walls are not distinctly
striated. Plate 2, figs. 20, 21, and 22, show three different
portions of the transverse section of the pericarp. In fig. 20,
the sclerenchymatous tissue found in the region corresponding
to the rounded elevation or rib is indicated and the stone cells
of the middle portion of the stony region are seen tangentially
elongated. Figure 21 shows a portion close to the dorsal region
of the coccus, and fig. 22 illustrates the arrangement of the stone
cells, which are found on the flattened sides of the pericarp.
The seed coat measures about 0.65 millimeter in thickness.
In the outer part it is limited by a single layer of thin-walled
empty parenchyma cells with a more or less rectangular out-
line. Their outer walls are wavy and brownish in color. In the
surface view these cells appear polygonal in outline with wavy
cell walls. Sometimes they are slightly elongated and their
walls vary from four to six. Plate 2, fig. 24, shows a transverse
section of the seed coat while fig. 25 on the same plate illustrates
the characteristics of the surface view of the outer layer of
cells of the seed coat. The inner part of the seed coat consists
mostly of obliterated or considerably flattened parenchyma cells.
Owing to the compactness of these cells their individual char-
acteristic cannot be determined. These flattened cells are dif-
ferentiated into two groups or regions. The walls of the cells
in the outer region are brownish in color, while those in the
inner region are colorless and hyaline. These two regions are
sometimes separated by a row of tangentially elongated paren-
chyma cells with thin walls. Boat-shaped structures filled with
protoplasm are often observed among the obliterated paren-
chyma cells in the innermost part of the seed coat.
The cotyledons in transverse section are plano-convex in out-
line. They are built up entirely of thin-walled parenchyma cells
filled with aleurone grains or protein granules. In the outer
part, they are surrounded by single thin-walled parenchyma
cells of rectangular, barrel-shaped or polygonal outlines. The
peripheral portion is occupied largely by radially elongated
thin-walled parenchyma cells, and the middle portion by poly-
gonal parenchyma cells. Plate 3, fig. 28, represents a segment
taken from the convex or dorsal side of a cotyledon, showing
the slightly elongated parenchyma cells. Figure 18, on same
plate, illustrates a segment taken from the ventral or flat side
of a cotyledon. This portion shows a greater elongation in the
parenchyma on the inner side. In the tangential section, these
46,2 Santos: Philippine Coriaria 265
parenchyma cells appear more or less isodiametric in charac-
teristic and they are polygonal in outline, as indicated on Plate
3, fig. 27. Plate 2, fig. 26, represents a median section of a
cotyledon cut parallel to its flat surface showing a group of
small elongated parenchyma cells, which initiate the forma-
tion of the conducting tissue.
When the seeds are subjected to Schultz's maceration process,
the conspicuous type of cell observed under the microscope is
indicated on Plate 2, fig. 23, a and 6. The stone cells derived
from the stony region display a great diversity of forms and
sizes. They vary from 0.01 to 0.06 millimeter in length. They
are either elliptical ovate, oblong, elongated, tapering at both
ends, straight or bent at one side, crooked or irregularly shaped
cells. Their walls are prominently pitted but not distinctly
striated and their cavities are very much reduced. The long
sclerenchymatous elements are numerous, and they are either
straight or crooked' and taper at both ends.
SUMMARY
1. The leaf in cross section is bifacial. The upper and lower
epidermis consist of a single layer of cells with very thick and
highly cutinized outer cell walls. The mesophyll is thin and
composed of two rows of palisade chlorenchyma in the upper
part and spongy chlorenchyma in the lower part.
2. The stomata are located on the lower surface only and are
characterized by the parallel arrangement of the two small
subsidiary cells to the pore. Epidermal outgrowth and calcium
oxalate are wanting.
3. In the surface preparation, both the upper and the lower
epidermal cells are polygonal in outline with fine wavy stria-
tions.
4. The midrib in cross section exhibits poorly developed collen-
chyma and sclerenchymatous cells. The endodermis is distinct
and the phloem cells are found surrounding the water conducting
tissue.
5. The seed is campylotropous, exalbuminous and inclosed
by the adhering pericarp. Externally, it is brown to dark brown
in color and characterized by the presence of riblike structure
elevations found at the dorsal and lateral sides.
6. The pericarp is hard and differentiated into two regions;
namely, the parenchymatous region and the stony region. The
seed coat is only one and very thin.
266 The Philippine Journal of Science
7. The embryo is small, slightly bent, and whitish in color.
The hypocotil is short and conical. The cotyledons are plano-
convex in outline, consisting of thin-walled parenchyma cells
richly supplied with protein granules and the plumule is incon-
spicuous.
BIBLIOGRAPHY
1. Kariyonb, T., and T. Sato. Coriaria japonica A Gray. Chem. Zentralbl.
101, 1 (1930) 3317.
2. Lindsay, W. L. On the toot-poison of New Zealand. Journ. Bot. 1
(1863) 247-50.
3. Matsumura, J. Notes on Liuku and Formosan plants. Bot. Mag. 12
(1898) 62.
4. Merrill, E. D. New or noteworthy Philippine plants, V. Philip. Journ.
Sci. 1 (Supplement) (1906) 205-6.
5. Reutter, L. Traite Matiere Medicale (1923) 275-6.
6. Solereder, H. Solereder's Systematic Anatomy of the Dicotyledons,
(Engl. Ed. Boodle and Fritsch) 1 (1908) 249.
7. Wells, A. H. The physiological active constituents of certain Philip-
pine medicinal plants, III. Philip. Journ. Sci. 14 (1919) 1-7.
ILLUSTRATIONS
[The microscopical drawings and sketches of some of the flowers were prepared by the
author, except figs. 20 and 30 which were drawn by Miss Teodora Kalalo, assistant
instructor, Department of Botany, University of the Philippines. The habit sketches,
including the sketches of some of the flowers, fruit, and seeds, are by Mr. Macario
Ligaya, Bureau of Science.]
Plate 1. Coriaria intermedia Matsumura
Fig. 1. A small portion of the plant showing the arrangement and char-
acter of the leaves and fruit; X 0.5.
2. A raceme of young bisexual flowers showing the bracts and
their characteristic filamentous stigma; X 2.5.
3. A portion of a reeeme of young male or sterile flowers; X 3.
4. A single bisexual flower after fertilization;, X 8.
5. A young bisexual flower partially dissected to show the relative
position between sepals s, petals p, the pistil and the stamen
drawn from a fresh flower; x 8.
6. A single bisexual flower showing the relative positions of the sta-
mens and stigmas; X 8.
7. A fully developed male flower with ten stamens with long filaments,
s, sepals, and p, petals;, X 8.
8. A longitudinal floral diagram of a young bisexual flower showing
the relative position between the sepals, petals, and pistils.
9. A young male or sterile flower showing the arrangement of the
anthers; X 8.
10. A semi-diagrammatic drawing of a mature bisexual flower, s,
sepals, p, petals.
11. A floral diagram of the cross section of a sterile or male flower.
12. A floral diagram of the cross section of a bisexual or perfect
flower.
13. A semi- diagrammatic drawing of a cross section of a nearly ma-
ture fruit, s, sepals, and p, petals.
14. A mature fruit; X 2.5.
15. a-e, A group of anthers dissected from a bisexual flower; X 12.
16. Normal stamens from a male flower, a, ventral view and 6, dor-
sal view; X 12.
17. The mature coccus drawn in three positions, a, dorsal side, b,
lateral side, and c, ventral side; X 6.
18. A diagrammatic representation of the transverse section of a
coccus, scl, sclerenchyma, p, parenchyma, and co, cotyledon and
h, xylem; X 16.
19. A lateral view of the embryo with one cotyledon removed, r,
radicle, and co, cotyledon; X 16.
267
268 The Philippine Journal of Science
Plate 2. Coriaria intermedia Matsumura
Fig. 20. A segment of the transverse section of the pericarp taken from
the lateral part near the region of a rib. scl, sclerenchyma,
sc, stone cells, and p, parenchyma ;. X 275.
21. Another segment of a transverse section from the lateral part of
the pericarp, e, epidermis, p, parenchyma, scl, sclerenchyma,
sc, stone cells; X 275.
22. A segment of the transverse section near the dorsal side of the
pericarp showing the greatly tangentially arranged scleren-
chyma cells, scl; and the stone cells, sc; X275.
23. A group of isolated cells from the pericarp, a, sclerenchyma cells,
X 165; and b, stone cells; X 450.
24. A segment of a transverse section of the seed coat, op, outer
obliterated parenchyma with brown coloration. op2, obliter-
ated parenchyma without brown colorations; X 275.
25. A segment of the surface view of the outer part of the seed
coat; X 275.
26. A longitudinal section of the cotyledon showing the elongated
parenchyma cells, which initiate the development of the con-
ducting tissue; X 275.
Plate 3. Coriaria intermedia Matsumura
Fig. 27. A tangential section of the cotyledon showing the polygonal out-
line of the parenchyma cells containing protein granules; X 275.
28. A segment of the transverse section of the cotyledon through the
dorsal side showing the epidermis and the slight radial elon-
gation of the parenchyma cells containing protein granules;
X275.
29. A transverse section of the cotyledon through the ventral side
showing the radial elongation of the parenchyma cells con-
taining protein granules; X 275.
30. A transverse section of a leaf through the midrib, c, collenchy-
ma, en, endodermis, scl, sclerenchyma, ph, phloem, and x, xy-
lem; X 125.
31. A segment of the surface preparation of the upper epidermis
showing a fine striation; X 450.
32. A surface view of a segment prepared from the lower epidermis
showing the stomata and the characteristic fine striation of
the cuticle; X 450.
Plate 4. Coriaria intermedia Matsumura
Fig. 33, &-c. Photograph of the portions of the fresh leaflless branches
showing the arrangement of unisexual and bisexual flowers.
Santos: Philippine Cortaria.]
[Philip. Journ. Sci., 46, No. 2.
PLATE 1.
Santos: Philippine Coriaria.]
[Philip. Journ. Scl, 46, No. 2.
PLATE 2.
Santos : Philippine Coriaria.]
[Philip. Journ. Sci., 46, No. 2.
PLATE 3.
Santos: Philippine Coriaria.]
[Philip. Journ. Scl, 46, No. 2.
PLATE 4.
NEW OR LITTLE-KNOWN TIPULIDJE FROM THE
PHILIPPINES (DIPTERA), XI1
By Chaeles P. Alexander
Of Amherst, Massachusetts
THREE PLATES
The very interesting crane flies discussed herewith were taken
in various parts of Luzon by my friends Messrs. McGregor,
Duyag, and Rivera, and in Minadanao by my former student at
this college Mr. Charles F. Clagg. I wish to thank these gentle-
men for their continued kindly interest in making known this
fauna.
TIPULIN^E
Genus DOLICHOPEZA Curtis
Dolichopeza Curtis, Brit. Entomol. (1825) 62.
I must consider several groups that are allied to Dolichopeza
and have hitherto been maintained as distinct genera as repre-
senting no more than subgeneric aggregations. Such subgenera
are as follows:
Dolichopeza, s. s., is found in the western Patearctic and
eastern Nearctic Regions, with the vast majority of the species
occurring in Australia and New Zealand. Curiously enough,
with the above distribution, no species is found in the Chilean
Subregion of the Nootropics. A few aberrant species that
may be found to be incorrectly placed herein, including isolata
Alexander (Luzon), are found in the Oriental and Ethiopian
Regions.
Nesopeza Alexander is the dominant subgenus in the Oriental
and eastern Palsearctic Regions. The typical group (gracilis
and allies) has Rs very long and the wings handsomely pat-
terned. Edwards is inclined to restrict the subgeneric name to
this latter group, leaving the equally or more abundant species
with plain wings and Rs of a shorter length in the typical
subgenus.
Contribution from the entomological laboratory, Massachusetts Agri-
cultural College.
269
270 The Philippine Journal of Science 1931
Mitopeza Edwards includes five species, of which one is de-
scribed herewith from Luzon. All the known species are from
the Oriental Region. The typical form, having macrotrichia in
the distal cells of the wing, with very blunt unchitinized cerci,
and with a very remarkable condition of the spermothecal ducts
in the female, is very distinct. However, longicornis Brunetti
lacks the macrotrichia and has the spermothecal ducts reduced
in number and size. This reduction is carried still further in
rizalensis sp. nov. The Bornean mjobergi Edwards has a very
distinct and peculiar venation (Plate 1, fig. 4).
Oropeza Needham has about a score of species that are nearly
evenly distributed in the eastern Nearctic and eastern Palsearctic
Regions.
Trichodolichopeza Alexander, as known, is entirely Ethiopian,
with the majority of the species occurring in South Africa.
Megistomastix Alexander is represented only by two species
in the Greater Antilles of the Neotropical Region.
These subgeneric groups may be separated by means of the
following key:
1. Apical cells of wings with macrotrichia 2.
Apical cells of wing without macrotrichia 4.
2. Cell 1st M2 closed 3.
Cell 1st M2 open by atrophy of basal section of M3.
Trichodolichopeza Alexander.
3. Rs short, transverse, not exceeding one-half R2+3; Sc2 ending opposite
origin of Rs;; m-cu connecting with M3+4 at or close to fork, always
beyond the fork of M Megistomastix Alexander.
Rs long, exceeding one-half R2+3, Sc2 nearly opposite its fork; m-cu
connecting with M far before its fork, at the fork in nigromaculata
Edwards Mitopeza Edwards (in part) (nitidirostris et al.).
4. Cell 1st M2 closed 5.
Cell 1st M2 open by atrophy of basal section of M3 6.
5. Rs short, Sc2 ending opposite or just beyond its origin; m-cu beyond
fork of M Oropeza Needham.
Rs long, subequal to or nearly as long as R2+3 Sc2 ending nearly op-
posite its fork; m-cu far before fork of M.
Mitopeza Edwards (in part) (longicornis).
6. Rs short, transverse, Sc2 opposite or close to its origin.
Dolichopeza Curtis.
Rs longer, subequal to or longer than R2+8, Sc2 ending beyond mid-
length 7.
7. Branches of medial field regularly pectinate, r-m connecting posteriorly
with the basal section of M1+2 (Plate 1, fig. 3).... Nesopeza Alexander.
Branches of medial field not pectinate, r-m connecting with Mi some
distance beyond origin (Plate 1, fig. 4).
Mitopeza Edwards (in part) (mjobergi) .
46.2 Alexander: Philippine Tipulidse, XI 271
Scamboneura Osten Sacken might also be construed as fall-
ing within the limits of Dolichopeza, but I would believe that it
represents a separate branch of the Dolichopezaria.
DOLICHOPEZA (MITOPEZA) RIZALENSIS sp. nov. Plate 1, fig. 1; Plate 2, fiff. 23.
General coloration dark brown ; legs with the tips of the tibiae
and all tarsi snowy white; wings grayish subhyaline, with a
heavy dark brown pattern in the costal and apical portions;
sparse macrotrichia in cells of wing at apex.
Male. — Length, about 8 millimeters ; wing, 10.5 ; antenna, about
4.5.
Female. — Length, about 9 millimeters; wing, 10 to 11.
Rostrum brownish yellow; palpi black. Antennae (male) a
little more than one-half the length of body; basal segments
testaceous, beyond the first flagellar passing into black; flagellar
segments long-cylindrical, with a delicate erect pubescence and
a group of three or four relatively short verticils at base on
outer face of segments, these much shorter than the segments
alone. Head blackish, sparsely pruinose behind on sides, the
front yellowish.
Mesonotal prsescutum dark brown, with indications of four
dark reddish brown stripes; posterior sclerites of mesonotum
more uniformly brown. Pleura testaceous brown. Halteres
elongate, dark brown, the extreme base of stem pale yellow.
Legs with the coxse and trochanters testaceous yellow; femora
dark brown, paler basally ; tibiae brown at base, the tips narrowly
snowy white; tarsi snowy white. In the male the tibiae are
chiefly white, the basal third more darkened. Wings (Plate 1,
fig. 1) grayish subhyaline, heavily patterned in costal and apical
portions with dark brown ; cells C and Sc dark, the bases paler ;
radial field heavily darkened, especially in female, with conspi-
cuous whitish spots before and beyond the stigma; cord and
veins beyond it seamed with brown. Sparse macrotrichia in
outer ends of cells R3 to 2d M2, inclusive. Venation: Rs sub-
equal to or longer than R2+3; cell 1st M2 relatively small.
Abdominal segments chiefly blackened, especially on posterior
portion, the base laterally brightening to obscure yellow; hypo-
pygium dark. Male hypopygium (Plate 2, fig. 23) with the ter-
gite, 9t, trifid, the pale cushionlike median lobe projecting caudad
beyond the level of the laterals, densely clothed with microscopic
erect setulse ; lateral lobes with less numerous coarse setse. Outer
dististlyle, od, profoundly bifid. Inner dististyle, id, very irreg-
ular in outline.
262412-
272 The Philippine Journal of Science 1931
Ovipositor with blunt fleshy lobes; spermothecal ducts rel-
atively few.
Luzon, Rizal Province, Novaliches, August 8 and 9, 1930 (A.
C. Duyag) ; holotype, male; allotype, female; paratypes, 1 male
and 1 female.
Dolichopeza (Mitopeza) rizalensis agrees with the subgeno-
type, D. (M.) nitidirostris (Edwards), and the more recently
described D. (M.) nigromaculata (Edwards) in the presence of
macrotrichia of the apical cells of the wing, differing from both
in the snowy- white tarsi and tibial apices. In the latter char-
acter, the present species agrees with D. (M.) longicornis (Bru-
netti), which differs in having no macrotrichia in apical cells of
wing and with the male antennae longer than the entire body.
The following key will suffice to separate the known species of
Mitopeza:
1. Cell 1st M8 open by the atrophy of the basal section of M» (Borneo).
mjobergi (Edwards) .
Cell 1st M* closed 2.
2. Apical cells of wing without macrotrichia (Assam).
longicornis ( Brunetti ) .
Distinct though sparse macrotrichia in apical cells of wing 3.
3. Legs black 4.
Legs with the tarsi and tips of tibiae extensively whitened (Luzon).
rizalensis sp. nov.
4. Head blackish; prsescutum with four velvety-black spots, one pair in
humeral region, the other before the wings; m-cu at fork of M
(Perak) nigromaculata (Edwards).
Head dark brown; thorax dark brown, the prsescutum with indications of
three darker stripes; m-cu far before fork of M (Kedah).
nitidirostris ( E d wards ) .
DOLICHOPEZA (NESOPEZA) MELANOSTERNA sp. nov. Plate 1, fig. 2; Plate 2, fi*. 24.
Male. — Length, about 8 to 9 millimeters; wing, 9.5 to 11.
Female. — Length, about 11 millimeters; wing, 11.
Generally similar and allied to D. (N.) angustaxillaris Alex-
ander (Luzon), differing especially in the larger size, details
of venation, as the deeper forks of M, the much darker colora-
tion, including the entirely blackened eighth sternite, and the
structure of the male hypopygium, especially the ninth tergite.
Antennae longer than in angustaxillaris, the flagellar segments
correspondingly lengthened. Mesonotum dark brown, the pleura
pale, the dorsopleural region darkened; ventral sternopleurite,
meron, and anepisternum with darkened areas. Legs with the
white proximal ends of fore basitarsi narrow, of the mid basi-
tarsi obsolete or with a mere genual brightening. Wings
46,2 Alexander: Philippine TipuUdse, XI 273
(Plate 1, fig. 2) strongly tinged with blackish; medial forks deep.
Abdominal tergites chiefly blackened, with a narrow transverse
yellow annulus on basal half; hypopygium black, including the
entire eighth sternite. Male hypopygium (Plate 2, fig. 24) with
the tergite, 9t, blackened, the lateral lobes broad, simple, their
ventral margins heavily blackened and microscopically toothed;
median lobe small, acute. Outer dististyle, od, shorter and
stouter than in angustaxillaris.
Luzon, Laguna Province, Ube, February 3 to 12, 1930 (F.
Rivera) ; holotype, male; allotype, female; paratypes, 8 males.
LIMONIIN^E
LIMONIINI
LIMONIA (LIMONIA) BILOBULIFERA sp. nov. Plate 1, fig. 5; Plate 2, fig. 25.
General coloration pale ocherous, with a conspicuous black dor-
sal stripe on pleura; knobs of halteres infuscated; legs chiefly
yellow, the femoral tips insensibly darkened ; wings whitish sub-
hyaline, the small stigma circular in outline; cell 1st M2 long;
male hypopygium with the outer dististyle a small, hairy, bilobed
structure; inner style with the body small, produced into a long
ribbonlike portion.
Male. — Length, about 4.2 to 4.4 millimeters ; wing, 5 to 5.3.
Rostrum reddish brown ; palpi a little darker. Antennae black,
the outer segments paling to brown; relatively elongate for a
member of this genus, if bent backward extending nearly to the
wing root; flagellar segments passing through oval to cylin-
drical, the verticils of the outer segments shorter than the seg-
ments alone; terminal segment elongate, about one-half longer
than the penultimate. Head blackish, sparsely pruinose; eyes
(male) contiguous on vertex or nearly so, the ommatidia coarse.
Mesonotum pale ocherous, scarcely variegated with darker,
the scutellum and median area of scutum more testaceous.
Pleura pale yellow, with a broad blackish dorsolongitudinal
stripe. Halteres pale, the knobs infuscated. Legs with the
coxae and trochanters yellow ; femora yellow, the tips insensibly
darker; tibiae and tarsi obscure yellow, the terminal tarsal seg-
ments infuscated; claws elongate, with a long conspicuous spine
at near one-third the length. Wings (Plate 1, fig. 5) whitish
subhyaline, the prearcular and costal regions more yellowish;
stigma brown, circular; veins dark brown, paler in the flavous
areas. Venation: Scx ending about opposite two-thirds the
length of the long arcuated Rs, Sc2 at its tip ; free tip of Sc2 and
R2 in transverse alignment ; cell 1st M2 unusually long, the second
274 The Philippine Journal of Science 1931
section of Mi+2 being equal to the third section ; basal section of
Ms longer than m, gently arcuated; m-cu at fork of M; vein 2d
A at origin converging toward 1st A, the cell relatively long and
narrow.
Abdominal segments bicolorous, dark brown, the caudal mar-
gins of the individual segments narrowly obscure yellow ; hypo-
pygium chiefly yellow, the basistyles conspicuously dark brown.
Male hypopygium (Plate 2, fig. 25) with the basistyles, 6, covered
with short dense setulse, in addition to scattered major setse;
ventromesal lobe, h, large, flattened, basal in position, weakly
bilobed at apex. Two dististyles, the outer, dd, a small, unequal-
ly bilobed hairy structure ; inner style, vd, with the base a trifle
enlarged, thence long-produced into a slender blade, the inner
margin before midlength with a small pale spinous point. Gon-
apophyses, g, with the mesal-apical lobe elongate, slender, trans-
versely ribbed.
Luzon, Laguna Province, Ube, February 3, 1930 (F. Rivera) ;
holotype, male; paratype, male.
Limonia (Limonia) bilobulifera is very different from the
other regional species in the structure of the male hypopygium.
LIMONIA (LIMONIA) MELANOPLEURA sp. noy. Plate 1, fi«. 6; Plate 2, fig. 26.
General coloration brownish black, including most of the tho-
racic pleura; halteres and legs brownish black; claws simple;
wings with a strong blackish tinge, the circular stigma darker;
Sc2 ending about opposite midlength Rs, Sc2 at its tip; male
hypopygium with the ventromesal lobe of basistyle very large
and stout; ventral dististyle small, setiferous, produced into a
long slender rostral prolongation, without spines.
Male. — Length, about 3.5 millimeters; wing, 4.2.
Female. — Length, about 5 millimeters; wing, 4.8.
Rostrum and palpi black. Antennae black throughout; fla-
gellar segments (male) oval, the longest verticils slightly exceed-
ing the segments and unilaterally arranged ; the female has the
segments short-oval. Head black, sparsely pruinose; eyes of
both sexes contiguous or nearly so, at most separated by a
capillary strip of anterior vertex.
Mesonotum brownish black. Pleura chiefly black, the ven-
tral sternopleurite and dorsopleural region paler, testaceous
brown. Halteres brownish black, the extreme base of stem pale.
Legs with the coxae brownish testaceous, the trochanters some-
what paler; remainder of legs brownish black; claws small,
without distinct spines. Wings (Plate 1, fig. 6) with a strong
46>2 Alexander: Philippine Tipulidse, XI 275
blackish tinge, the circular stigma darker brown; veins dark
brown. Venation: Scx ending about opposite midlength of Rs,
Sc2 at its tip; free tip of Sc2 slightly proximad of R2; m-eu
just before the fork of M; vein 2d A long, converging toward
1st A at origin.
Abdomen, including the hypopygium, black. Male hypopy-
gium (Plate 2, fig. 26) with the tergite, 9t, unusually extensive,
broad at base, strongly narrowed outwardly, the two low lobes
separated by a small emargination ; a submarginal series of about
six strong setae on either side. Basistyle, b, relatively long and
slender, the ventromesal lobe very stout, occupying almost the
entire mesal face of the style. Ventral dististyle, vd, a small
oval lobe with long conspicuous setae, the rostral prolongation
long, slender, only gently curved, with no developed spines.
Luzon, Laguna Province, Ube, April 14, 1930 (R. C. McGre-
gor) ; holotype, male ; allotype, female.
Limonia (Limonia) melanopleura is well-distinguished by the
small size, very extensive black coloration, and the structure of
the male hypopygium. I cannot indicate any closely allied re-
gional species.
LIMONIA (LIMONIA) TREMULA sd. nov. Plate 1, fi&. 7.
General coloration of mesonotal prsescutum brown, variegated
with sublateral and a posterior median yellow stripe; pleura
yellow, with a conspicuous longitudinal dark stripe; halteres
dusky; legs yellow; wings grayish yellow, with a restricted
brown pattern; Rs angulated and weakly spurred at origin;
m-cu about one-half its length beyond the fork of M ; abdomen
brownish black, the segments narrowly ringed caudally with
yellow.
Female. — Length, about 5.5 millimeters; wing, 6.3.
Mouth parts small, the rostrum reddish brown; palpi black.
Antennae black throughout ; basal flagellar segments subglobular
to short-oval, the outer segments more elongate; segments with
two conspicuous verticils on outer face, unilaterally arranged.
Head fulvous brown, the center of the vertex extensively dark-
ened.
Pronotum brown. Mesonotal praescutum brown, variegated
with brownish yellow, the latter including sublateral stripes that
meet in front and a median stripe on posterior half of sclerite;
the darkened portions include the lateral margins to the anterior
region and submedian stripes that become approximated in front,
behind crossing the suture onto the scutal lobes ; median region
276 The Philippine Journal of Science 1931
of scutum and scutellum obscure yellow, the latter with each lat-
eral third darkened; postnotal mediotergite testaceous brown,
more yellowish laterally and on the dorsal half of the pleuroter-
gite. Pleura obscure yellow, with a conspicuous dorsolongitudi-
nal dark stripe that extends to the abdomen, including the ven-
tral half of the pleurotergite ; ventral sternopleurite a little
darkened. Halteres dusky, the base of stem restrictedly pale.
Legs with the coxse yellow, the fore coxae a trifle darkened;
trochanters yellow ; remainder of legs obscure yellow, the femoral
tips rather broadly but insensibly clearer yellow; terminal tar-
sal segments a trifle darkened; claws relatively slender, with a
basal tooth that is further prolonged into a slender seta. Wings
(Plate 1, fig. 7) grayish yellow, with a restricted and relatively
diffuse brown pattern, including the stigma, cord, and outer end
of cell 1st M2 ; origin of Rs and fork of Sc ; basal portion of wing
and costal region a little darkened ; veins cream-colored, a little
darkened in the clouded areas. Macrotrichia of veins relatively
long and conspicuous, including Rs except on its basal section.
Venation : Scx ending about opposite three-fifths the length of Rs,
Sc2 at its tip; Rs angulated and weakly spurred at origin; free
tip of Sc2 and R2 in approximate transverse alignment; cell 1st
M2 large, subrectangular, a little longer than vein M3 beyond it ;
m-cu about one-half its length beyond the fork of M, subequal
to the distal section of C\xt ; vein 2d A strongly sinuous, at origin
parallel to vein 1st A or nearly so.
Abdomen brownish black, the segments narrowly ringed cau-
dally with yellow; genital segments ocherous. Ovipositor with
the tergal valves slender, gently upcurved, reddish horn color;
sternal valves large, straight, conspicuously blackened at base.
Luzon, Laguna Province, Ube, February, 1930 (F. Rivera) ;
holotype, female.
Limonia (Limonia) tremula is amply distinct from described
regional species, agreeing in some respects with L. (L.) luteivit-
tata Alexander, but differing in all details of coloration and
venation.
LIMONIA (LIBNOTES) UNISTRIOLATA sp. nov. Plate 1, fi&. 8; Plate 2, fi*. 27.
General coloration of mesonotal prsescutum obscure yellow,
with a single complete black stripe, on either side behind bor-
dered by clear golden yellow; rostrum, palpi, antennae, knobs of
halteres and legs black; wings with a faint brown suffusion; Scx
long; Rs angulated at origin; cell 1st M2 small, rectangular, less
than one-half the distal section of M1+2; anal veins gently diver-
46,2 Alexander: Philippine Tipulidx, XI 277
gent; male hypopygium with the ventral dististyle large and
fleshy, the rostral prolongation with two very unequal spines.
Male. — Length, about 6 millimeters; wing, 6.8.
Female. — Length, about 6.3 millimeters; wing, 7.
Rostrum and palpi black. Antennae black throughout; fla-
gellar segments subcylindrical, becoming longer outwardly; ver-
ticils of basal segments slightly exceeding the segments; ter-
minal segment pointed at apex, about one-half longer than the
penultimate. Head black, sparsely pruinose, the anterior vertex
more silvery, reduced to a narrow strip.
Pronotum black, the anterior lateral pretergites yellow. Me-
sonotal praescutum obscure yellow, with a single broad and
complete black stripe, on either side on posterior two-thirds
clear golden yellow; lateral portions of sclerite weakly in-
fumed; scutal lobes black; median region of scutum paler; scutel-
lum and postnotal mediotergite blackened. Pleura chiefly brown,
the posterior dorsopleural region and the ventral sternopleurite
obscure yellow. Halteres pale, the knobs dark brown. Legs
with the fore coxae inf uscated, the remaining coxae and all tro-
chanters yellow ; remainder of legs dark brown, only the femoral
bases restrictedly pale. Wings (Plate 1, fig. 8) with a faint
brown suffusion, the circular stigma a trifle darker; veins pale
brown. Venation: Scx ending opposite r-m, Sc2 far from its
tip, Sc2 about opposite midlength of the angulated Rs; free tip
of Sc2 and R2 in transverse alignment; cell 1st M2 relatively
small, less than half the distal section of vein M1+2; m and
basal section of M3 subequal, straight, in approximate trans-
verse alignment; m-cu at one-third the length of cell 1st M2;
anal veins parallel to gently divergent at origin.
Abdomen dark brown, the sternites obscure yellow; hypopy-
gium dark. Male hypopygium (Plate 2, fig. 27) with the tergite,
9t, extensive, the caudal margin with a deep V-shaped notch,
the lateral lobes with coarse setae. Basistyle, &, relatively small.
Ventral dististyle, vd, a large fleshy lobe, the rostral prolongation
with two unequal gently curved spines. Dorsal dististyle a
strongly curved hook, the tip acute. Gonapophyses, g, with
the concave margin of the mesal-apical lobe with minute points.
Luzon, Mountain Province, Ifugao Subprovince, Huungduan,
April 5, 1930 (F. Rivera) ; holotype, male; allotype, female.
Allied to species such as L. (L.) neofamiliaris Alexander and
L. (L.) subfamiliaris Alexander, likewise from Luzon, differing
conspicuously in the coloration and details of structure of the
male hypopygium.
278 The Philippine Journal of Science 1931
LIMONIA (LIBNOTES) MELANCHOLICA sp. nov. Plate 1, fig. 9; Plate 2, fig. 28.
General coloration polished black; rostrum, palpi, antennae,
knobs of halteres, and legs blackened ; wings with a faint dusky
tinge, cells C and Sc darker; S^ ending some distance beyond
r-m, Sc2 opposite the fork of Rs; cell 1st M2 rectangular, less
than one-half vein M1+2 beyond it; m and basal section of M3
in nearly transverse alignment ; m-cu at about one-fourth to one-
fifth the length of cell 1st M2; anal veins gently divergent; male
hypopygium with the ventral dististyle large and fleshy, the ros-
tral prolongation with two straight spines of unequal diameter.
Male. — Length, about 5.5 to 7 millimeters; wing, 6 to 7.
Female. — Length, about 6.5 to 6.8 millimeters ; wing, 6.5.
Rostrum, palpi, and antennae black; flagellar segments oval,
becoming more elongate outwardly; longest verticils exceeding
the segments in length and unilaterally arranged. Head black,
heavily dark gray pruinose; anterior vertex narrow, light gray.
Pronotum black. Mesonotum polished black, the median re-
gion of scutum obscure yellow. Pleura chiefly black, the pro-
pleura, dorsal pteropleurite, and dorsopleural membrane brown-
ish yellow. Halteres yellow, the knobs dark brown. Legs with
the fore and hind coxae yellow, the mid-coxae slightly darkened ;
all trochanters yellow; remainder of legs brownish black, the
femoral bases restrictedly pale ; claws relatively long and slender,
with an acute subbasal tooth, with additional microscopic basal
denticles. Wings (Plate 1, fig. 9) with a faint dusky tinge, cells
C and Sc more inf umed ; wing tip and posterior margin to vein
Cu slightly clouded; a dark seam along vein Cu; stigma sub-
circular in outline, slightly darker than the ground color; veins
dark brown. Venation: Sc^, ending some distance beyond r-m,
Sc2 opposite the fork of Rs, Scx a little longer than m-cu ; free tip
of Sc2 and R2 in approximate transverse alignment; cell 1st M2
rectangular, less than one-half vein M1+2 beyond it; m and basal
section of M3 in nearly transverse alignment ; m-cu at from one-
fourth to one-fifth the length of cell 1st M2 ; anal veins gently
divergent.
Abdomen black, the sternites brown; genitalia of both sexes
darkened. Male hypopygium (Plate 2, fig. 28) with the caudal
emargination of the tergite, 9t, broadly V-shaped'; marginal setae
of lobes strong and powerful; a group of about three small me-
dian setae. Ventral dististyle, vd, large and fleshy, the rostral
prolongation with two nearly straight spines, of nearly equal
length but unequal diameter, the inner slender to setif orm ; setae
46,2 Alexander: Philippine Tipulidse, XI 279
of apex of prolongation relatively sparse. Dorsal dististyle a
chitinized sickle, sinuously to subangularly bent, the long acute
tip slightly decurved. Gonapophyses, g, with the apex of each
slightly blackened, the surface and margin before tip with erect
pale points.
Luzon, Tayabas Province, Candelaria, near town, alongside a
small stream, June 20 to 25, 1930 {McGregor and Rivera) ; holo-
type, male ; allotype, female ; paratypes, 5 of both sexes.
Limonia (Libnotes) melancholica is allied to L. (L.) neofa-
miliaris Alexander and L. (L.) subfamiliaris Alexander, to-
gether with the species described herewith as L. (L.) unistrio-
lata sp. nov., differing in the almost uniformly black color, in
addition to details of the venation and male hypopygium.
LIMONIA (LIBNOTES) PERRARA sp. nov. Plate 1, fig. 10.
General coloration of prsescutum yellow in front, with four
brown stripes behind; pleura yellow, with two black longitudi-
nal stripes; halteres yellow; legs yellow, the femora with a
broad dark brown subterminal ring; wings yellow, handsomely
patterned with brown; Rs only slightly arcuated; m-cu just
before midlength of cell 1st M2.
Male. — Length, about 7.5 millimeters; wing, 8.5.
Female. — Length, about 7.5 millimeters; wing, 9.
Rostrum and palpi ocherous, the latter narrowly darkened
at tips. Antennae with the scape pale, the flagellum somewhat
darker; flagellar segments short-oval to subcylindrical, crowd-
ed, gradually increasing in size outwardly, the terminal seg-
ment long; verticils relatively short and inconspicuous, not or
scarcely exceeding the segments in either sex. Eyes of male
large, contiguous; of female separated for a long distance only
by a capillary strip of vertex; posterior portion of head gray.
Mesonotal prsescutum in front chiefly yellow, more saturated
anteriorly; four pale brown stripes on posterior half; scutal
lobes brown, the median portion, with adjoining parts of prse-
scutum and scutellum, whitish ; caudal margin of scutellum nar-
rowly blackened on either side; postnotal mediotergite gray,
with a capillary pale median line. Pleura yellow, with two
conspicuous blackish longitudinal areas, including a narrow dor-
sal stripe from the propleura to the abdomen, the second area
including almost all of the sternopleurite. Halteres pale yellow.
Legs with the coxae and trochanters pale yellow ; femora yellow,
with a broad dark brown subterminal ring, the apical yellow
280 The Philippine Journal of Science 1931
portion very narrow; tibiae yellow; basal segments of tarsi yel-
low, the terminal three and distal end of the second blackened;
claws with a conspicuous spine at near one-third the length,
with additional smaller spines nearer the base. Wings (Plate
1, fig. 10) creamy yellow, with a handsome brown pattern, in-
cluding four areas in cell Sc, the first two not encroaching on
cell C, the second sending a triangular cloud along Rs; fourth
area including R2 and tip of Sc2 ; cord and outer end of cell 1st
M2 seamed with brown; a broad seam on R2+3 for almost the
entire length; a series of five oval clouds on distal portions of
veins R3 to M4, inclusive, placed shortly before the margin;
posterior margin of wing almost to tip narrowly clouded with
brown; brown clouds at ends of veins Cn1 and 2d A, the latter
extended basad for about one-half the length of the vein ; axilla
darkened ; veins yellow, brown in the clouded areas. Venation :
Scx ending just before the proximal end of m, Sc2 at its tip;
free tip of Sc2 and the spur of R1+2 subequal, or the latter
greatly reduced so that R2 and the free tip of Sc2 are in approx-
imate transverse alignment; R2 unusually long; Rs gently
curved ; m nearly twice as long as the basal section of M3, gently
arcuated; m-cu just before midlength of cell 1st M2; cell 1st
A at margin very much wider than cell Cu; anal veins at base
almost parallel, thence divergent.
Abdomen dark brown, the caudal margins of the tergites in
the male yellow ; sternites paler ; in female, the abdomen is more
uniformly yellow, variegated laterally with blackish areas. Ovi-
positor dark, the cerci weakly bidentate at tips, there being a
small dorsal subterminal denticle, as in the group.
Luzon, Mountain Province, Benguet Subprovince, Pauai, alti-
tude 8,000 feet, April 21 and 22, 1930 (F. Rivera) ; holotype,
male; allotype, female.
Limonia (Lihnotes) perrara is a member of a group of the
subgenus that includes L. (L.) amatrix Alexander (Japan), L.
(L.) klossi Alexander (Federated Malay States), L. (L.) terraz-
reginse Alexander (Queensland), and possibly other species, in
wihich the ovipositor has the cerci distinctly toothed on dorsal
margin before apex. The nearest relative of the present species
appears to be amatrix, which differs in veitational details, as the
very strongly arcuated Rs, the position of m-cu at about one-
fourth the length of cell 1st M2, and other details, and in the very
distinct leg pattern.
46,2 Alexander: Philippine Tipvlidm, XI 281
LIMONIA (DICRANOMYIA) ORTHIA sp. nor- Plate 1, fi*. 11; Plate 2, figr. 29.
General coloration dark brown; rostrum and antennse black;
halteres pale; wings milky white, with a heavy dark pattern
that is chiefly marginal in distribution, there being a series of
four darker costal areas, with gray clouds at wing tip and at
ends of anal veins; Sc2 far from tip of Scx; male hypopygium
with the spines of the rostral prolongation short, placed close
together on the small prolongation.
Male. — Length, about 5 millimeters; wing, 5.5.
Rostrum, palpi, and antennse black throughout. Head brown-
ish gray; anterior vertex narrow.
Mesonotum dark brown, the scutellum somewhat paler.
Pleura blackish, pruinose. Halteres pale. Legs with the coxse
brownish testaceous, the fore coxse darker; trochanters testa-
ceous; remainder of legs pale brown; claws with a single long
basal spine. Wings (Plate 1, fig. 11) with the ground color
milky white, the prearcular and costal regions more yellowish;
a heavy brown pattern that is chiefly marginal in distribution,
including a series of four areas along the costal margin, the
first being at areulus, the second at Sc2, the third at end of Sct
and origin of Rs, the last stigmal ; wing tip in outer end of cell
R3 clouded with gray ; large gray clouds at ends of anal veins ;
cord and outer end of cell 1st M2 seamed with gray; veins pale,
darker in the clouded areas, yellow in the brightened costal por-
tions. Venation : Scx ending opposite origin of Rs, Sc2 far from
its tip, at near midlength of vein R ; m-cu close to fork of M ;
cell 2d A moderately wide.
Abdomen dark, the incisures paler ; male hypopygium with the
basistyles dark, the large ventral dististyles paler. Male hypo-
pygium (Plate 2, fig. 29) with the basistyles, b, small, the ven-
tromesal lobe large. Dorsal dististyle a very strongly curved
pale sickle, the extreme tip upcurved. Ventral dististyle, vd,
a large fleshy lobe, the rostral prolongation small, the two
spines straight, subequal in length and size, about as long as the
prolongation itself. Gonapophyses, g, with the mesal-apical
lobe gently curved to the acute tip.
Luzon, Mountain Province, Benguet Subprovince, Pauai, alti-
tude 8,000 feet, April 21, 1930 (F. Rivera) ; holotype, male.
Limonia (Dicranomyia) orthia is allied to the larger Jap-
anese species, L. (D.) mesosternata (Alexander) and L. (D.)
mesosternatoides (Alexander), differing very conspicuously in
the structure of the male hypopygium.
282 The Philippine Journal of Science 1931
LIMONIA (DICRANOMYIA) NEOPUNCTULATA sp. nov. Plate 1, fl*. 12; Plate 2, fig. 30.
Male.— Length, about 4.5 millimeters; wing, 4.8.
Generally similar and allied to L. (D.) punctulata, differing
especially in the details of structure of the male hypopygium.
The general coloration, wing pattern, and venation (Plate 1, fig.
12) are quite the same in both species.
Male hypopygium (Plate 2, fig. 30) with the dorsal dististyle,
dd, subangularly bent beyond midlength. Ventral dististyle,
vd, relatively small, the rostral prolongation with a single short
stout spine from a raised tubercle, the spine about equal in
length to the prolongation, evidently formed by the coalescence
of two spines, the suture being evident. Gonapophyses, g, with
the mesal apical lobe simply bifid.
Limonia (D.) punctulata (de Meijere) is well distinguished by
the details of the hypopygium (Plate 2, fig. 31), especially the
very slightly curved dorsal dististyle, the long, very slender ros-
tral spine, vd, that is strongly curved at tip and without a basal
tubercle, and the irregularly toothed gonapophyses, g.
Limonia (D.) fullowayi (Alexander) has the male hypopy-
gium (Plate 2, fig. 32) with the dorsal dististyle, dd, very
strongly curved to an acute point ; rostral spine, vd, single, long,
and very slender, without basal tubercle, entirely straight ; gona-
pophyses, g, not evidently bifid at tips.
Mindanao, Davao district, Calian, Lawa, May 3, 1930, at
light (C F. Clagg) ; holotype, male.
It is very evident that several species of Limonia center about
punctulata in the Oriental-Eastern Palsearctic faunal regions.
The three species compared above, having a single spine on
the rostral prolongation of the ventral dististyle, and with the
gonapophyses variously toothed at apices, seem to be well-sepa-
rated by the genitalic differences as described. Limonia (D.)
subpunctulata Alexander (Formosa) is distinct in the bispinous
rostral prolongation. Limonia (D.) fascipennis (Brunetti),
described from northern India, is possibly distinct from any of
the above. It was described from a single broken female and
since the name fascipennis has been used in Limonia (Limno-
bia) on two previous occasions, the name should be dropped until
the species is rediscovered at or near the type locality.
HELIUS (EURHAMPHIDIA) FUSCOFEMORATUS sp. nov. Plate 1, fig. 13.
Unusually large (wing, female, over 6.5 millimeters) ; rostrum
relatively elongate, about one -half longer than the remainder of
head; mesonotum dark brown, restrictedly paler laterally; legs
46,2 Alexander: Philippine Tipulidaz, XI 283
black, the tips of the tibiae narrowly snowy white, this including
about the distal sixth or less of the segment.
Female. — Length, about 7 to 7.5 millimeters; wing, 6.5 to 7.
Rostrum unusually long for a member of this subgenus, about
one-half longer than the remainder of head, black; palpi black.
Antennae with the basal segment obscure yellow beneath, the
remainder of the organ black ; flagellar segments oval, with ver-
ticils that exceed the segments. Head brownish gray.
Pronotum dark brown, restrictedly yellow behind. Mesono-
tum chiefly dark brown, the lateral portions of praescutum paler.
Pleura brownish yellow. Halteres dusky, the base of the stem
restrictedly yellow. Legs with the coxae and trochanters yellow;
femora brownish black, with no sign of brightening at genua;
tibiae black, the tips narrowly snowy white, on the posterior legs
this including less than the distal sixth ; tarsi white, the terminal
segments blackened. Wings (Plate 1, fig. 13) with a pale
brownish tinge, the oval stigma darker brown; prearcular and
costal regions slightly more yellowish; a yellowish seam in cell
M adjoining vein Cu; veins brown. Venation: S^ ending op-
posite r-m, Sc2 at its tip; basal section of M1+2 subequal to
second section, the inner end of cell 1st M2 being pointed ; m-cu
before midlength of cell 1st M2.
Abdominal tergites dark brown, the sternites yellow, the sub-
terminal segments more darkened. Ovipositor with the tergal
valves slender, brownish black, their acutely upcurved tips pale.
Luzon, Mountain Province, Benguet Subprovince, Pauai, alti-
tude 8,000 feet, April 26, 1930 (F. Rivera) ; holotype, female;
paratype, female.
Helius (Eurhamphidia) fuscofemoratus may be confused only
with H. (E.) nigrofemoratus (Alexander), which differs con-
spicuously in the small size, the short rostrum, and the in-
creased amount of white on apices of tibiae.
HELIUS (EURHAMPHIDIA) INDIVISUS sp. nov. Plate 1, fisr. 14; Plate 2, figr. 33.
Male. — Length, about 4.6 millimeters; wing, 5.4.
Similar to H. (E.) diacanthus (Alexander) and H. (E.) ab-
normalis (Brunetti) in general appearance, differing especially
in the structure of the male hypopygium.
Rostrum pale brown, a little longer than the remainder of
head ; palpi dark brown. Antennae black. Head dark gray, the
narrow anterior vertex more silvery gray.
Thoracic dorsum reddish brown, the median area of praescu-
tum a little darker. Pleura more testaceous yellow, the dorsal re-
284 The Philippine Journal of Science 1931
gion a little darker. Halteres pale, the knobs dusky. Legs with
the coxae pale ; femora brown, the tips broadly and conspicuously
white, tibiae brown, the bases narrowly white, the amount about
one-third that of the femoral tips; tibial tips broadly snowy
white; tarsi white, the terminal segments darkened. Wings
(Plate 1, fig. 14) whitish subhyaline, the stigmal region darker;
veins pale brown. Venation: Scx ending about opposite r-m,
Sc2 at its tip; m short to very short, cell 2d M2 narrowed at base;
cell 1st M2 short, subquadrangular, m-cu at near midlength.
Abdominal tergites light brown, the sternites pale yellow.
Male hypopygium (Plate 2, fig. 33) with the lateral spines of
the tergite, 9i, simple, not bearing basal spinules or lobes, as in
diaeanthus and abnormalis. Outer dististyle, od, with delicate
but distinct erect setae for almost the whole length ; in diaeanthus
and abnormalis the style is glabrous. Inner dististyle, id, nar-
row, terminating in two larger setae, the margin at base almost
smooth, not expanded and provided with conspicuous tubercles,
as in diaeanthus.
Mindanao, Davao district, Calian, La Lun Mountains, altitude
5,800 feet, July 3, 1930 (C. F. Clagg) ; holotype, male.
ERIOPTERINI
TRENTEPOHLIA (MONGOMA) DISTALIS sp. nov. Plate 1, fig. 15.
General coloration dark brown to brownish black; antennae
black throughout ; halteres dusky ; legs black, the terminal tarsal
segments paling to brownish yellow; wings with a dusky tinge,
the costal region more blackened ; R3 not conspicuously arcuated ;
cell 1st M2 small, m-cu beyond the fork of M ; abdominal tergites
black.
Male. — Length, about 7 millimeters ; wing, 7.2.
Female. — Length, about 7 millimeters; wing, 7.2.
Rostrum and palpi dark, the tips of the labial palpi pale yellow.
Antennae black; flagellar segments long-oval to subcylindrical,
with elongated verticils. Head black, the anterior vertex re-
duced to a linear strip.
Mesonotum dark brown to brownish black, the median region
(of scutum and lateral portions of scutellum somewhat paler.
Pleura dark yellowish brown, the propleura and dorsopleural
membrane dark brown. Halteres dusky. Legs with the fore
coxae dark brown, the remaining coxae and all trochanters more
testaceous brown; remainder of legs black, the terminal tarsal
segments paling to brownish yellow; legs without specially devel-
46,2 Alexander: Philippine Tipvlidde, XI 285
oped armature of any kind. Wings (Plate 1, fig. 15) with a
dusky tinge, cells C and Sc more blackish; the small ill-delimited
stigma and an apical suffusion paler brown; veins brownish
black. Venation: Sc± ending just beyond proximal end of R2;
Rs shorter than R2+3+4; R2 at or close to the fork of R3+4; R3
gently sinuous but not conspicuously arcuated at origin ; cell 1st
M2 small, the fusion of R4+5 and Mi+2 subequal to or one-half
longer than the second section of M1+2> the proximal end of cell
R5 lying proximad of any other beyond the cell ; m-cu from two-
thirds to nearly its own length beyond the fork of M, at beyond
one-third the length of cell 1st M2 ; apical fusion of C^ and 1st
A slight.
Abdominal tergites and hypopygium black; basal sternites yel-
low, blackened laterally, beyond the second segment passing into
black. In the female, the sternites more uniformly brown, with
narrow glabrous apical margins. Ovipositor with the cerci rel-
atively long and slender.
Mindanao, Davao district, Calian, La Lun Mountains, altitude
5,500 feet, July 2 and 3, 1930 (C. F. Clagg) ; holotype, male; allo-
type, female. "Among ferns and on mossy trees." — Clagg.
By my most recent key to the Philippine species of Trente-
pohlia 2 the present species runs to couplet 8, disagreeing with
both included species in the position of m-cu, and in other char-
acters. It may be noted that there is a slight typographical
error in this couplet, the last symbol, M3, of the second alternative
(riverai) correctly being Mi+2 to agree with the first alternative
(brevifusa).
GONOMYIA (PROGONOMYIA) TEREBRELLA sp. nov. Plate 1, figr. 16.
General coloration of mesonotum reddish brown; antennae
black throughout; halteres dusky; wings with a strong brown
suffusion ; Sc long, Scx ending opposite the fork of Rs ; vein R3
at margin close to R1+2, cell R2 being very narrow ; ovipositor
with the tergal valves long and chitinized, the sternal valves
reduced to tiny blackened hairy lobes.
Female. — Length, about 5 millimeters ; wing, 4.8.
Rostrum and palpi black. Antennae black throughout, rel-
atively elongate for this sex ; flagellar segments oval, the verticils
exceeding the segments. Head brown, sparsely pruinose.
Pronotum whitish. Mesonotal praescutum reddish brown, the
median area blackened ; scutum reddish brown, the centers of the
* Philip. Journ. Sci. 43 (1930) 297-298.
286 The Philippine Journal of Science 1931
lobes conspicuously blackened ; scutellum dark medially at base,
the apex broadly testaceous ; postnotal mediotergite black, sparse-
ly pruinose. Pleura dark brown, variegated with light and
dark areas, the obscure yellow including the dorsopleural mem-
brane and areas dorsad of the mid- and hind-coxss ; the blackened
areas occur as spots on the dorsal anepisternum and dorsal pter-
opleurite. Halteres dusky. Legs with the coxse infuscated;
trochanters obscure yellow ; remainder of legs black, the femoral
bases broadly obscure yellow. Wings (Plate 1, fig. 16) with
a strong brown tinge, the stigmal region vaguely and diffusely
darker; veins brownish black. Macrotrichia of costa and veins
relatively long and conspicuous. Venation : Sc long, Scx ending
opposite the fork of Rs, Sc2 some distance from its tip, Scx
being equal to B2+3+4; R3 and R4 strongly divergent, R3 at margin
closely approaching R1+2, cell R3 being very narrow at margin ;
cell 2d M2 deep; m-cu a short distance beyond the fork of M,
in alignment with the other elements of the cord.
Abdomen black, the subterminal sternites paler; genital
sheaths blackened. Tergal valves of ovipositor elongate,
straight, reddish horn color; sternal valves reduced to tiny
blackened hairy lobes, directed ventrad.
Mindanao, Davao district, Calian, June 13, 1930, trap lantern
set at edge of forest (C. F. Clagg) ; holotype, female.
Gonomyia (Progonomyia) terebrella is closest to G. (P.) tene-
brosa Edwards (Siam) in the general coloration and structure
of the ovipositor, differing in the details of venation, especially
the unusually long Sc, which ends opposite the fork of Rs. The
fly differs more widely from G. (P.) brunnescens Edwards
(Borneo) in coloration and venation.
Genus ERIOPTERA Meigen
Subgenus TELENEURA subgen. nor.
Characters as in the typical subgenus, differing especially in
the wing venation. Mesonotal prsescutum with longitudinal
rows of long erect setse on interspaces. Veins and cells beyond
the cord very elongate, the cord lying at or before midlength of
the wing (Plate 1, fig. 17). Rs very short, subequal to or only
a little longer than R2+3+4; cell 1st M2 open by atrophy of m;
m-cu at fork of M; veins M4 and Cux deflected only slightly,
cephalad at their tips ; vein 2d A only gently sinuous.
In typical Erioptera, Rs is three or more times as long as
R2+3+4, the slightly oblique cord lying at or beyond three-fifths
the length of the wing; vein 2d A very strongly sinuous, the
distal third or fourth paralleling the anal margin of wing.
46'2 Alexander: Philippine Tipvli&x, XI 287
Type of subgenus, Erioptera fusca de Meijere (Oriental
Region).
Other species pertaining to Teleneura are Erioptera argenti-
frons Edwards, E. melanotamia sp. nov., E. nigribasis Edwards,
E. parallels Brunetti, E. punctipennis Brunetti, and E. subfusca
Edwards, all Oriental. These species may be separated by means
of the following key:
1. Wings variegated with dark areas, either on the membrane itself or as
conspicuous darkened hair patches on the veins 2.
Wings uniform in color 3
2. Femora yellow, the tips imperceptibly darkened (British India: Hima-
layas) punctipennis Brunetti.
Femora yellow, with about the basal half blackened (Malay Peninsula
and Borneo) nigribasis Edwards.
3. General coloration brownish ocherous, without conspicuous markings
(British India: Himalayas; Malay Peninsula) parallela Brunetti.
General coloration dark brown to black; if pale, variegated with black
longitudinal markings 4#
4. Halteres with at least the knobs yellow 5.
Halteres with the knobs blackened 6.
5. Halteres yellow; general coloration of thorax dark brown; male hypopy-
gium without conspicuous modified setae at apex of basistyle; gonapo-
physes simple, crook-shaped (Sumatra and Borneo).
subfusca Edwards.
Halteres with the stem black, the knobs yellow; general coloration of
thorax black; male hypopygium with a group of about five powerful
setae at apex of basistyle; gonapophyses bispinous, tonglike (Fed-
erated Malay States) argentifrons Edwards.
6. Thorax brown, the lateral margins of praescutum pale; dorsal thoracic
pleura with a narrow blackened longitudinal stripe (Luzon and Min-
danao) melanotmnia sp. nov.
General coloration of thorax uniform dark brown or black (Feder-
ated Malay States and Mindanao) fusca de Meijere.
ERIOPTERA (TELENEURA) FUSCA de Meijere.
Erioptera fusca de Meijere, Tijdsch. v. Entom. 56 (1913) 351.
La Lun Mountains, Calian, Davao district, Mindanao, altitude
5,800 feet, July 3, 1930 (C. F. Clagg). The specimens are
almost black, instead of dark brown, but there seems to be no
doubt as to the identity.
ERIOPTERA (TELENEURA) MELANOTiENIA sp. nov. Plate 1, fi*. 17; Plate 2, fiff. 34.
Mesonotal praescutum light brown, margined with obscure
yellow; pleura pale, with a black dorsolongitudinal stripe; knobs
of halteres brownish black ; wings with a brown tinge.
Male. — Length, about 2.5 millimeters ; wing, 3.
Female. — Length, about 3 millimeters ; wing, 3 to 3.2.
262412 9
288 The Philippine Journal of Science 1931
Rostrum and palpi black. Antennae black. Head light ocher-
ous, dark brown in center of vertex and on occiput.
Mesonotal praescutum and scutum brown to dark brown, the
lateral margins paling to obscure yellow. Pleura obscure yel-
low, including the dorsopleural region and dorsal pleurotergite
lying above, and the dorsal meron and sternopleurite lying below,
a broad black dorsal stripe that extends from the propleura to
the abdomen ; ventral sternopleurite and meron again darkened.
Halteres obscure yellow, the knobs brownish black. Legs with
the fore coxae dark, the remaining coxae and trochanters obscure
yellow; femora obscure yellow, this coloration obscured by dark
setae. Wings (Plate 1, fig. 17) with a brownish tinge, the base
and costal region somewhat more yellowish brown; veins pale
brown, the macrotrichia dark. Venation : As in the subgenus ;
vein 2d A ending opposite m-cu.
Abdomen brownish black, the hypopygium paler. Male hypo-
pygium, (Plate 2, fig. 34) with the tergal plate (9£, one-half
figured) margined with conspicuous spines. Apex of basistyle,
b, without specially modified setae. Outer dististyle, od, pale,
at apex expanded into a blackened setiferous head; inner dis-
tistyle, id, a pale flattened blade, the distal third more nar-
rowed. Longest gonapophysis, g, more or less crook-shaped,
its apex cultriform, the two together appearing somewhat lyri-
form; shorter gonapophysis, g, more foot-shaped, the surface
with abundant delicate setae, including a tuft of longer setae at
the region of the "heel."
Luzon, Laguna Province, Ube, February, 1930 (F. Rivera) ;
holotype, male; allotype, female; numerous paratypes of both
sexes. Mindanao, Davao district, Calian, La Lun Mountains,
altitude 5,500 feet, July 2 to 5, 1930 (C. F. Clagg) ; paratypes,
5 males and females.
ERIOPTERA (EMPEDA) LUNENSIS sp. nov. Plate 1, fi*. 18; Plate 3, fi*. 35.
General coloration of praescutum brown medially, the lateral
portions gray; antennae black, the first flagellar segment pale
yellow; head blue-gray; halteres pale yellow; legs light brown,
appearing darker by a covering of scales and setae; Scx ending
opposite origin of Rs; cell R3 very deep.
Male. — Length, about 2.5 millimeters; wing, 2.8 to 3.
Female. — Length, about 3.5 millimeters ; wing, 3.3.
Rostrum and palpi black. Antennae with the scape and fla-
gellum black, the first flagellar segment abruptly pale yellow.
Head light blue-gray.
46,2 Alexander: Philippine Tipnlidse, XI 289
Pronotum and anterior lateral pretergites whitish. Mesonotal
prsescutum dark brown medially, the sides light gray to blue-
gray ; posterior sclerites of mesonotum chiefly darkened, the pos-
terior margin of scutellum more brightened, the postnotal medio-
tergite light gray pruinose. Pleura dark, sparsely pruinose,
the dorsopleural membrane restrictedly pale. Halteres pale
yellow. Legs with the coxae reddish brown, the fore coxae
darker; trochanters reddish brown; remainder of legs light
brown, the terminal tarsal segments passing into black; legs
with flattened scales, in addition to the usual setae. Wings
(Plate 1, fig. 18) grayish, the base and costal region more yel-
lowish; veins brown. Costal fringe relatively long and con-
spicuous. Venation : Sc-l ending opposite origin of Rs, Sc2 faint,
at tip of SCi ; R2 slightly oblique in position, shorter than R2+s+4
and about one-third R ; cell R3 unusually deep, approaching
the condition in typical Erioptera, vein R3 subequal to or only a
little shorter than Rs; cell M2 open; m-cu at or just before the
fork of M.
Abdominal tergites brown, the sternites paler ; hypopygium ob-
scure yellow. Male hypopygium (Plate 3, fig. 36) with the outer
dististyle, od, profoundly bifid, entirely glabrous, both arms
flattened and obtuse at tips. Inner dististyle, id, a pale flattened
blade, the distal half with microscopic sensory setse.
Mindanao, Davao district, Calian, La Lun Mountains, alti-
tude 5,500 to 5,800 feet, July 3 and 4, 1930 (C. F. Clagg);
holotype, male; allotype, female; paratypes, 12 of both sexes.
"Swept from among ferns and undergrowth on margins of
small brook ; others at trap lantern hung among ferns in dense
mossy forest." — Clagg.
Erioptera (Empeda) lunensis is most closely allied to E. (E.)
gracilis (de Meijere), differing in the coloration and in details
of venation, as the shorter Sc and deeper fork of cell R3. Both
species have conspicuous flattened and striated scales on the
legs, interspersed with the usual setse.
MOLOPHILUS BANAHAOENSIS sp. nov. Plate 1, fig. 19; Plate 3, fig. 36.
Belongs to the gracilis group; allied to M. kempi; antennae
(male) elongate; general coloration of body, antennae, halteres,
and legs blackish; wings tinged with blackish; vein 2d A rela-
tively short; male hypopygium with the dorsal lobe of basistyle
expanded at apex into a glabrous spatulate head ; ventral lobe of
basistyle with long coarse retrorse setse.
290 The Philippine Journal of Science tsai
Male. — Length, about 2.8 millimeters; wing, 3.5; antenna,
about 2.5.
Female. — Length, about 3 millimeters; wing, 3.2.
Rostrum and palpi black. Antennae with the scapal segments
obscure yellow; flagellum black; antennae (male) nearly as long
as entire body; flagellar segments fusiform, the apical necks
longer and slenderer than the narrow basal portion. Head
black, sparsely pruinose.
Mesonotum black, the humeral region restrictedly paler ; pseu-
dosutural foveae black; anterior lateral pretergites restrictedly
obscure yellow. Pleura black, the ventral sternopleurite and
meron a little paler. Halteres blackened, the base of the stem
obscure yellow. Legs with the coxae and trochanters obscure
yellow; remainder of legs blackened. Wings (Plate 1, fig. 19)
with a strong blackish tinge, the veins more seamed with darker,
the extreme wing tip pale ; veins and macrotrichia dark brown
to black. Venation: R2 and r-m in transverse alignment; vein
2d A relatively short, ending some distance before the proximal
end of m-cu.
Abdomen, including hypopygium, black. Male hypopygium
(Plate 3, fig. 36) with the basistyle, b, produced at apex into
four distinct lobes, the outermost a small glabrous spine on
outer dorsal margin; immediately laterad of this, on dorsal
margin a long hairy fingerlike lobe, the apex, db, expanded into
an obtuse glabrous spatula; mesal lobe flattened, narrowed out-
wardly and here provided with several long coarse setae ; ventral
lobe, vb, longer than the mesal, more or less clavate, at apex
with a group of very long, coarse, retrorse setae (only the bases
of which are shown in the figure), the longest about two-fifths
the entire lobe. Outer dististyle, od, a glabrous blackened spine,
the tip acute. Inner dististyle, id, subequal in length, yellow,
dilated on basal half, the inner margin on basal fifth with a few
setae; apex narrowed into a spine, with a few microscopic spi-
nulas on outer margin before apex.
Luzon, Laguna Province, Ube (R. C. McGregor) ; holotype,
male, February 12, 1930; allotype, female, April 14, 1930.
Molophilus banahaoensis is closely allied to M. kempi Alexan-
der (British India: Eastern Himalayas), differing especially in
the structure of the male hypopygium.
MOLOPHILUS PROCERICORNIS so. nov. Plate 1, fig. 20; Plate 3, fig. 37.
Belongs to the gracilis group; general coloration of mesono-
tum dark brown ; antennae (male) elongate ; pleura reddish yel-
low, variegated with brown; knobs of halteres weakly infus-
46,2 Alexander: Philippine Tipulidse, XI 291
cated ; male hypopygium large and conspicuous, the dorsal lobe
of the basistyle terminating in a flattened glabrous blade; two
dististyles, one an acutely pointed black spine.
Male. — Length, about 3.5 millimeters; wing, 4; antenna, about
2.8.
Rostrum reddish brown; palpi black. Antennae (male)
elongate, if bent backward extending to beyond midlength of
the body; scapal segments obscure yellow; flagellum black;
flagellar segments elongate-fusiform, with long outspreading
black verticils at thickest part. Head light gray, the anterior
vertex paler.
Anterior lateral pretergites pale yellow. Mesonotal praescu-
tum with the humeral and lateral portions pale yellow, the re-
mainder of disk chiefly covered by three dark brown stripes
that are confluent or nearly so; median vitta slightly divided
behind; scutal lobes dark brown; scutellum pale; postnotal me-
diotergite reddish brown. Pleura reddish yellow, variegated
with dark brown or dorsopleural membrane and anepisternum ;
ventral sternopleurite and meron darkened. Halteres pale, the
knobs weakly infuscated. Legs with the coxae and trochanters
yellow; remainder of legs obscure yellow, the vestiture chiefly
dark; tarsal segments passing into brown. Wings (Plate 1,
fig. 20) grayish yellow, the prearcular and costal regions brighter
yellow; veins brownish yellow, the macrotrichia a little darker.
Venation : R2 lying distad of the level of r-m ; vein 2d A relatively
short, ending before the caudal end of m-cu.
Abdominal tergites brownish black, the sternites paler, the
large hypopygium obscure yellow, with blackened dististyles.
Male hypopygium (Plate 3, fig. 37) with the basistyles, 6, rela-
tively short and stout; dorsal lobe, db, long and relatively slender,
setif erous for almost the entire length, the apex a short, sinuous,
glabrous blade; ventral lobe, vb, short and broad, with abundant
long retrorse setae; an additional ventral lobe (not figured),
small and very slender, pale, fleshy, with from six to eight setae
at and near apex, the total length being somewhat less than
the main ventral lobe. Two dististyles, the outer, od, blackened,
from a dilated flask-shaped base, the remainder a sinuous black
spine. Inner dististyle, id, a little longer, the basal two-thirds
or slightly more pale yellow, the gently curved apex blackened.
iEdeagus elongate, with a conspicuous lateral flange.
Mindanao, Davao district, Calian, La Lun Mountains, altitude
5,500 to 5,800 feet, July 2 to 4, 1930 (C. F. Clagg) ; holotype,
male; paratypes, 5 males.
292 The Philippine Journal of Science i9si
Molophilus procericornis is quite distinct from other mem-
bers of the gracilis group, the chief characters being antennal
and hypopygial. Females in the same collection do not seem
to be conspecific and are not further discussed.
MOLOPHILUS MENDICUS sp. nor. Plate 3, &g. 38.
Belongs to the gracilis group; general coloration of mesono-
tum brownish gray; antennae short in both sexes; halteres dusky;
wings pale grayish, the veins pale; vein 2d A relatively short;
male hypopygium with all lobes of basistyle fleshy and setif erous
to their obtuse tips, the outer lobe bearing a blackened spinous
point.
Male. — Length, about 2.8 millimeters; wing, 3.4.
Female. — Length, about 3.5 millimeters ; wing, 3.5.
Rostrum and palpi dark brown. Antennae short in both sexes,
brown throughout, in the female somewhat paler. Head gray-
ish brown.
Mesonotum brownish gray, the lateral margin and humeral
region somewhat brighter, inclosing the relatively small red-
dish brown pseudosutural foveae; scutellum obscure yellow,
darkened medially; postnotal mediotergite plumbeous brown.
Pleura plumbeous. Halteres dusky. Legs pale brown, the color
chiefly produced by the vestiture ; tips of tibiae and outer tarsal
segments darker. Wings with a pale grayish tinge, the veins
very pale; macrotrichia dark brown. Venation: R2 lying some
distance before the level of r-m, R2+3+4 thus shortened, about
two-thirds the basal section of R5; vein 2d A short, ending
before the level of the caudal end of m-cu.
Abdomen dark brown, the genitalia in both sexes more yel-
lowish. Male hypopygium (Plate 3, fig. 38) with the three
lobes of the basistyle, b, all fleshy, obtuse, provided with setae
to their tips; on margin of outer lobe, on inner face, a curved
blackened hook; mesal lobe small and slenderer. Two dis-
tistyles, d; these entirely pale and generally similar in outline,
one a little more expanded on basal half, the distal half slender,
with small subappressed spines before apex, at tip with two
or three setif erous punctures; second style a straight flattened
blade, slightly constricted at near midlength, at apex with a
very few weak spinous points. Phallosomic structure a pale
cushion that is densely set with microscopic setulae. iEdeagus
very long and slender, the base more dilated.
Mindanao, Davao district, Calian, La Lun Mountains, alti-
tude 5,800 feet, July 3, 1930 (C. F. Clagg) ; holotype, male; alio-
46,2 Alexander: Philippine Tipulidx, XI 293
type, female. "Among ferns and undergrowth along margins of
small brook." — Clagg.
Molophilus mendicus is a small, insignificant species that is
best characterized by the details of structure of the male hypo-
pygium.
MOLOPHILUS TAWAGENSIS sp. nov. Plate 3, fig. 39.
Belongs to the gracilis group; general coloration of mesono-
tum light grayish brown; antennae (male) short; pronotum and
anterior lateral pretergites white; pleura liver brown; halteres
with infuscated knobs; wings pale grayish, the costal region
clear light yellow, the posterior prearcular region infumed; male
hypopygium with only two lobes on the elongate basistyle, both
obtuse and with setae to their tips; a blunt setiferous lobe on
mesal face of basistyle near origin ; two dististyles.
Male. — Length, about 3.5 millimeters; wing, 4.5.
Rostrum and palpi black. Antennae (male) short, the basal
segments pale, the outer segments darkening to brown. Head
pale gray.
Pronotum and anterior lateral pretergites white. Mesonotal
praescutum light grayish brown, paling to clearer gray on sides ;
humeral and lateral portions pale yellow; pseudosutural foveae
relatively large, reddish brown ; median region of scutum gray-
ish; scutellum pale testaceous brown; postnotal mediotergite
gray. Pleura relatively dark liver brown. Halteres obscure
golden yellow, the knobs infuscated. Legs with the coxae brown-
ish yellow; trochanters obscure yellow; remainder of legs
chiefly brown, the terminal tarsal segments more blackened.
Wings pale grayish, the costal region clear light yellow; poste-
rior prearcular region darkened; veins pale, the macrotrichia
darker. Venation : R2 faint, lying opposite the basal section of
R5 ; vein 2d A ending just before the caudal end of m-cu.
Abdomen brown, the hypopygium yellow. Male hypopygium
(Plate 3, fig. 39) with the basistyles, b, relatively elongate, with
a very deep incision down the face, separating the lateral and
mesal lobes. Lateral lobes slender, much shorter than either
dististyle, with setae to the obtuse tip ; mesal lobe flattened ; on
mesal face of basistyle, closer to base, a broad lobe set with
coarse setae. Outer dististyle, vd, more slender, terminating
in a long acute spinous point, the surface at near midlength a
trifle roughened. Inner dististyle, id, broader, with a conspicu-
ous flange on basal half, the terminal bladelike portion with mi-
croscopic scattered setae. Surface of phallosomic structure with
delicate microscopic setulae.
294 The Philippine Journal of Science 1931
Luzon, Mountain Province, Ifugao Subprovince, Tawag, April
6, 1930 (F. Rivera) ; holotype, male.
Molophilus tawagensis belongs to the costalis subgroup, in-
cluding many species in the fauna of eastern Asia. The details
of the male hypopygium furnish the best characters for the sep-
aration of the various forms.
Genus STYRINGOMYIA Loew
Styringomyia Loew, Dipt. Beitr. 1 (1845) 6.
Idiophlebia GRUNBERG, Zool. Anzeiger 26 (1903) 524-528.
Pycnocrepis Enderlein, Zool. Jahrbucher 32 (1912) 65.
Mesomyites Cockerell, Proc. U. S. Nat. Mus. 52 (1917) 377.
The now rather numerous Philippine species of Styringomyia
may be separated in the male sex by means of the following key :
1. Wings with a strong blackish tinge, the basal fourth more yellowish;
legs black, the femora with a narrow yellow subterminal ring.
fumipennis Edwards.
Wings yellow or yellowish, immaculate, or spotted and washed with
darker 2.
2. General coloration of mesonotum gray; legs uniformly brown; wings
unmarked with darker; male hypopygium without specially enlarged
setae on apical lobe of basistyle mcgregori Alexander.
General coloration of mesonotum yellow, variegated with black; legs
yellow, the femora and tibiae ringed or spotted with brown; wings
yellow, patterned with brown, at least with a small darkened spot
at arculus; male hypopygium with the basistyle terminating in one
or two enlarged spinous setae 3.
3. Wings unmarked, except for a tiny darkened spot at arculus 4.
Wings spotted or washed on disk with darker , 5.
4. Mesonotum pale yellow, the praescutum without distinct markings; hal-
teres pale yellow; abdominal tergites with two small brown spots on
caudal margin; male hypopygium with the phallosome including a
flattened plate, its margin microscopically serrulate.
luteipennis sp. nov.
Mesonotum with the praescutum yellow, with two black lines before the
suture; halteres dusky; abdominal tergites with the marginal spots
confluent to form bands; male hypopygium not as above, the phal-
losome an elongate hook montina sp. nov.
5. Male hypopygium with basistyle at apex terminating in two spinous
setae 6.
Male hypopygium with basistyle at apex terminating in a single
setae 7.
6. Wings relatively long and narrow, the anterior branch of Rs subtrans-
verse; male hypopygium with the intermediate and inner arms of
dististyle small and inconspicuous armata Edwards.
Wings of normal shape, the anterior branch of Rs oblique, as usual in
the genus; male hypopygium with the inner arm of dististyle ex-
panded into an oval blade claggi sp. nov.
46,2 Alexander: Philippine Tipvlidse, XI 295
7. Male hypopygium with the outer arm of the dististyle a long slender
rod that terminates in a very long seta 8.
Male hypopygium with outer arm of the dististyle variously formed,
not bearing an apical seta.... 10.
8. Wings with the veins and cells behind the anterior margin strongly
washed with brown, the broad costal border yellow.
flavocostalis Alexander.
Wings yellow, with the usual four restricted dark clouds, located on
the anterior cord, outer end of cell 1st M2, m-cu, and distal end of
vein 2d A 9.
9. Wings with vein 2d A curved at end; male hypopygium with the ninth
sternite at apex very broad, heavily blackened, clothed with delicate
erect setae, the two enlarged apical bristles widely separated.
nigrosternata sp. nov.
Wings with vein 2d A short-spurred at end; male hypopygium with the
ninth sternite entirely pale, narrowed to a point outwardly, the
two apical bristles thus appearing approximated to actually contig-
uous ceylonica Edwards.
10. Male hypopygium with the outer arm of the dististyle a simple black-
ened spine, the tip acute tablasensis Alexander.
Male hypopygium with the outer arm of the dististyle a powerful struc-
ture, at apex produced mesad at a right angle into a spikelike point.
neocolona sp. nov.
STYRINGOMYIA FUMIPENNIS Edwards.
Styringomyia fumipennis Edwards, Notulae Entomologies 6 (1926)
37.
Type locality : Mount Banahao, Luzon. One male, Mount Ta-
buan, Cagayan, Luzon, May, 1929 (F. Rivera).
STYRINGOMYIA MCGREGORI Alexander.
Styringomyia megregori Alexander, Philip. Journ. Sci. 28 (1925)
373-374.
Type locality: Manila, October, 1924 (R. C. McGregor). Sev-
eral additional specimens, Manila, October, 1929 and 1930, at
light {McGregor) . Mr. Edwards informs me that he has seen
it from Borneo and the Andaman Islands.
STYRINGOMYIA LUTEIPENNIS sp. nov. Plate 3, fig. 40.
General coloration pale yellow, the mesonotal prsescutum
without distinct markings ; wings pale yellow, unmarked except
for a dusky spot at arculus ; halteres yellow ; abdominal tergites
with two separate brown spots on caudal margin of each ; male
hypopygium with a single lateral enlarged seta on basistyle;
dististyle expanded into a broadly flattened blade.
Male. — Length, about 5.5 millimeters; wing, 3.5 to 3.7.
Rostrum and palpi yellow. Antennal scape brownish yellow,
especially on lower face; flagellum entirely pale yellow. Head
pale yellow.
296 The Philippine Journal of Science iosi
Mesonotum pale yellow, the praescutum without distinct mark-
ings; postnotal mediotergite with narrow brown lateral lines.
Pleura yellow. Halteres pale yellow. Legs yellow, the femora
with two restricted brown areas on outer face only; tibiae with
an incomplete brown ring before midlength, the tips inf uscated ;
tarsi yellow, the last segment dark brown. Wings pale yellow,
unmarked except for a small dusky area at arculus ; veins deeper
yellow but still very indistinct. Venation: Anterior branch of
Rs oblique; cell 2d M2 short-sessile to more broadly sessile, in
rare cases with a very short petiole ; m-cu about its own length
beyond the fork of M ; vein 2d A curved gently to margin.
Abdomen yellow, each tergite with two brown spots on caudal
margin; hypopygium yellow. Male hypopygium (Plate 3, fig.
40) small, the ninth tergite, 9*, terminating in a cordate seti-
ferous lobe, the apex narrowed but obtuse. Ninth sternite,
9s, broad, pale, with two widely separated spinous setse, the
intervening space very gently concave. Basistyle, b, with a
single developed apical spinous seta, its basal lobe small; a
reduced set beside the major spine. Dististyle, d, with the
outer arm pale, terminating in the usual very long seta, at base
with a group of about fifteen spines and a marginal comb of ten
to twelve close-set spines ; main blade of dististyle broadly flat-
tened, with abundant long black spinous setae; two pale arms
at base of dististyle, the shorter with marginal setae, the outer-
most a stout black spine; longer cephalic arm slenderer, ter-
minating in a group of six or seven stout spines. Phallosome,
p, with a group of spinous setae on either side; a flattened dark
plate, its apex truncate, the margins microscopically serrulate.
Luzon, Laguna Province, Mount Maquiling, January 29, 1930
(A. C. Duyag) ; holotype, male; paratypes, 8 males; above Ube,
foot of Mount Banahao, February 3 to 6, 1930 (F, Rivera) ;
paratypes, 2 males.
Styringomyia luteipennis much resembles S. flava Brunetti
and S. taiwanensis Alexander in the yellow wings, but belongs
to a different section of the genus, having but a single spinous
seta at apex of basistyle of male hypopygium. The small brown
spot at arculus and the structure of the male hypopygium furnish
distinctive features.
STYRINGOMYIA MONTINA sp. nov. Plate 3, fit?. 41.
Generally similar and closely related to S. luteipermis sp. nov.,
differing in slight details of coloration and structure of the
male hypopygium. Size larger and form stouter. First scapal
46,2 Alexander: Philippine Tipvlidse, XI 297
segment beneath and entire second segment blackened. Meso-
notal praescutum with two blackish lines before the suture.
Halteres dusky. Wings somewhat deeper yellow, especially in
the radial field, the veins correspondingly more distinct. Ab-
dominal tergites with the margins on caudal margin large, con-
fluent, to form apical bands. Male hypopygium (Plate 3, fig,
41) generally as in luteipennis, but the phallosome, p, entirely
different, terminating in an elongate hook, on outer margin
with numerous erect spinous setae and true spines.
Luzon, Mountain Province, Ifugao Subprovince, Pakawan,
April 7, 1930 (F. Rivera); holotype, male; paratypes, 2 males;
Banaue, April 4, 1930 (F. Rivera) ; allotype, female.
STYRINGOMYIA ARMATA Edwards. Plate 3, fi*. 42.
Styringomym armata Edwards, Ann. & Mag. Nat. Hist. IX 13 (1924)
274; Treubia 9 (1927) 355, fig. b.
Type locality : Mindanao. Lawa, Calian, Davao district, Min-
danao, April 28, 1930 (C. F. Clagg) ; Calian, July 14, 1930 (C.
F. Clagg), at light of house. The latter specimen is accom-
panied by the following note : "This walked across table with a
sort of dancing motion, raising its body up and down, at regular
intervals of about one-half second."
I believe the identification to be correct, despite certain de-
tails lacking in the original description. The present fly has the
wing unusually long and narrow for a member of the genus,
with the anterior branch of Rs subtransverse, as shown (Plate
1, fig. 21). The male hypopygium (Plate 3, fig. 42) is again
illustrated, the chief characters being the bispinous basistyle,
6, and the great reduction in size of the intermediate and poste-
rior branches of the dististyle, d.
STYRINGOMYIA CLAGGI sp. nov. Plate 1, fig. 22; Plate 3, fig. 43.
General coloration yellow, heavily variegated with black ; palpi
and antennal scape black, the flagellum yellow; head and thorax
without flattened setae; legs with complete rings on femora
and tibiae; male hypopygium with two apical spinous setae on
basistyle, these arising from elongate tubercles; main arm of
dististyle a broadly flattened blade.
Male. — Length, about 6 to 6.3 millimeters; wing, 4.5 to 5.
Female. — Length, about 5 to 5.5 millimeters; wing, 4 to 4.5.
Rostrum and palpi brownish black. Antennae with the scape
black, the flagellum abruptly pale yellow, the outer segments a
trifle more darkened. Head blackish, without flattened setae.
298 The Philippine Journal of Science 1931
Pronotum obscure yellow medially, more blackened laterally.
Mesonotal praescutum with the disk obscure yellow, the margin
and two intermediate vittae before the suture more blackened;
scutum with the median area and centers of the lobes obscure
yellow, the latter margined with blackish ; scutellum blackened,
the median region restrictedly obscure yellow; postnotal medio-
tergite black, with a capillary yellow median vitta. Pleura
obscure yellow, the dorsal sclerites darker. Halteres obscure
yellow ; knobs dark brown. Legs with the coxae and trochanters
pale yellow; femora yellow, with two broad complete brownish
black rings, in addition to the narrowly darkened tips ; the more
basal yellow annulus a little wider than the inclosing dark rings ;
outer yellow annulus narrow; tibiae yellow, the tips and a sub-
equal ring on basal half black; tarsi yellow, the outer segment
blackened, the narrow tips of the other segments infuscated.
Wings (Plate 1, fig. 22) yellow, with ill-delimited brown washes,
including the anterior cord, vein Cu and vein 2d A; veins pale
brown, C, Sc, and R more yellowish. Venation : Anterior branch
of Rs normally oblique; cell 2d M2 sessile; vein 2d A curved
gently to the margin.
Abdominal tergites light brown, the caudal margins darker
brown, the sternites and hypopygium yellow. Male hypopygium
(Plate 3, fig. 43) with the apical lobe of the ninth tergite, 9t,
low and obtuse, densely hairy. Ninth sternite 9s, narrowed
apically, the terminal setae not widely separated. Basistyle, b,
with two relatively short apical spinous setae from long basal
tubercles. Outer arm of dististyle, d, a long pale structure with
the usual very elongate terminal seta; main arm of dististyle
a broadly flattened blade, with long setoid spines that are chiefly
marginal in distribution, there being a row along outer edge and
a dense patch on mesal margin at near midlength.
Mindanao, Davao district, Calian, La Lun Mountains, altitude
5,500 to 5,800 feet, July 3 and 4, 1930, by sweeping vegetation
(C F. Clagg) ; holotype, male; allotype, female; paratypes, 1
male, 1 female; Mount Apo, 7,000 to 8,000 feet, September 21,
1930 (Clagg) ; paratypes, 1 male and 1 female.
This interesting Styringomyia is dedicated to my friend Mr.
Charles F. Clagg, who has, collected very numerous new and
rare Tipulidae in the highest mountains of Colorado and Min-
danao. The species is allied to S. ensifera Edwards, S. armata
Edwards, and S. acuta Edwards, in the bispinous basistyle of
the male hypopygium, differing in the unmodified setae of the
46>2 Alexander: Philippine Tipulidse, XI 299
head and thorax, and the details of the hypopygium, notably
the greatly expanded inner arm of the dististyle.
STYRINGOMYIA FLAVOCOSTALIS Alexander.
Sty ring omy id flavocostalis Alexander, Philip. Journ. Sci. 27 (1925)
76-77.
Type locality: Mount Maquiling, Luzon. Additional speci-
mens, Ube, Laguna Province, Luzon, altitude 300 to 400 meters,
January 26, 1930 (R. C. McGregor).
STYRINGOMYIA NIGROSTERNATA sp. nov. Plate 3, fig. 44.
General coloration chiefly pale; rostrum, palpi, and antennal
scape blackened; mesonotal praescutum with a broad medial
darkening; pleura yellow; wings pale yellow, with the usual
dark spots; male hypopygium with the basistyle unispinous,
this spinous seta arising from a very long basal tubercle ; ninth
sternite heavily blackened at apex, clothed with delicate erect
black setae.
Male. — Length, about 6 millimeters; wing, 4.6.
Rostrum and palpi brownish black. Antennae with the scapal
segments black, the flagellar segments brownish yellow. Head
brownish gray, the usual setae stout but not flattened.
Pronotum gray medially, brownish black laterally. Mesonotal
praescutum chiefly ocherous, with a sparse gray bloom, the me-
dian region on anterior half with a broad brownish black stripe;
a small blackish spot on either side at the suture, this area ex-
tended across the suture and partially encircling the scutal lobes
on outer side; scutellum pale, with a dark spot on either side;
postnotal mediotergite chiefly dark brown. Pleura light yellow.
Halteres yellow, the knobs slightly more orange. Legs with the
coxae and trochanters pale yellow; femora yellow, with two nar-
row, incomplete brown rings; tibiae yellow, the tips and an in-
complete ring before midlength brown; tarsi yellow, the tips
of the individual segments weakly darkened. Wings pale yellow,
with the usual four or five brown clouds, these being on anterior
cord, union of M2 and M3, fork of M3+4, m-cu, and the distal third
of vein 2d A ; veins yellow, dark brown in the inf uscated areas.
Venation: Anterior branch of Rs normally oblique; cell 2d M2
short-petiolate ; vein 2d A curved strongly into the margin, not
angulated.
Abdominal tergites yellow, the caudal margins of the segments
with two small brown triangles, these becoming larger and con-
fluent on the outer segments ; in addition to the above, a median
300 The Philippine Journal of Science 1931
brown clouding on basal half of tergites, on outer segments
heavier and more clearly delimited; sternites and hypopygium
yellow. Male hypopygium (Plate 3, fig. 44) with the apical lobe
of the tergite, 9t, long-triangular, the tip obtuse. Ninth sternite,
9s, broad, the apex extensively and conspicuously blackened, the
two apical spines unusually short, arising from small elevated
tubercles, the surface of the lobe with short erect black setae.
Basistyle, b, with a single terminal spinous seta, this unusually
short, less than one-half the long basal tubercle. Dististyle, d,
with the outer arm terminating in a long seta, without spines
at base; intermediate arm produced laterad into a long acute
spine at near midlength, the base of this spine and the arm be-
yond with a row of black spine3 ; inner arm a curved chitinized
rod, the tip obliquely acute and slightly blackened ; outer margin
of arm at midlength with a linear group or crest of about ten to
twelve spines; mesal face of arm at base with a group of long
spinous setae.
Mindanao, Davao district, Lawa, at trap lantern, April 24,
1930 (C. F. Clagg) ; holotype, male.
Styringomyid nigrosternata is very different from other re-
gional species in the structure of the male hypopygium, especially
the dististyle and ninth sternite.
STYRINGOMYIA CEYLONICA Edwards.
Styringomyia ceylonica Edwards, Ann. & Mag. Nat. Hist. VIII 8
(1911) 62-63.
Type locality: Weligama, Ceylon. The following authentic
Philippine records are available: Badajoz, Tablas, August 28,
1928 (F. Rivera and A. C. Duyag) ; Lawa, Davao district, Min-
danao, at light, April 24, 1930 (C. F. Clagg) .
Bezzi 3 recorded this species from Los Banos and Mount Maqui-
ling, but this record is almost certainly erroneous, as previously
indicated by Edwards.4
STYRINGOMYIA TABLASENSIS Alexander.
Styringomyia tablasensis Alexander, Philip. Journ. Sci. 40 (1929)
344-345.
Type locality: Badajoz, Tablas, August 27, 1928 (F. Rivera
and A. C. Duyag). Other Philippine records: Lawa, Davao dis-
trict, Mindanao, at light, April 24, 1930 (C. F. Clagg) ; Calian,
Mindanao, July 12, 1930, at light (C. F. Clagg) .
8 Philip. Journ. Sci. § D 12 (1917) 115.
4 Notulae Entomologicae 6 (1926) 34.
46,2 Alexander: Philippine Tipulidse, XI 301
STYRINGOMYIA NEOCOLONA sp. iiov. Plate 3, fi*. 45.
Closely allied to colona; general coloration yellow, the prae-
scutum with black lines behind ; blackened areas on femora and
tibiae restricted in area; male hypopygium with the apical lobe
of the tergite truncate; ninth sternite expanded at apex and
deeply emarginate.
Male. — Length, about 6 millimeters ; wing, 4.3.
Rostrum brown; palpi brownish yellow, the outer segment
paling to yellow. Antennae with the basal segment black above,
the remainder of organ pale yellow. Head light brown.
Pronotum restrictedly pale medially, blackened laterally. Me-
sonotal praescutum obscure brownish yellow; marked with black
behind, including two submedian black lines that converge in
front, inclosing an oval ocherous median area before the suture;
scutal lobes similarly ocherous, bordered externally by black;
scutellum obscure yellow, margined caudally by black; postnotal
mediotergite black. Pleura yellow. Halteres yellow. Legs
with the coxae and trochanters pale yellow; femur yellow,
with two small black spots on upper surface only; tibiae yellow,
the tips blackened, with an additional restricted black cloud on
upper surface before midlength; tarsi yellow, the terminal seg-
ment blackened. Wings pale yellow, with four blackish areas,
as usual in the genus, these on anterior cord, m and adjoining
veins, m-cu, and on the distal two-fifths of vein 2d A ; veins yel-
low, blackened in the dark areas. Venation: Anterior branch
of Rs normally oblique; m short but present, cell 2d M2 being
short-sessile; vein 2d A curved into the anal margin, the cell
relatively wide.
Abdominal tergites yellow, the segments with two small brown
spots on caudal margin, those of the second segment large, of
segments three to five small, on the outer segments again be-
coming larger and confluent; sternites and hypopygium yellow.
Male hypopygium (Plate 3, fig. 45) with the apical lobe of the
tergite, 9t, elongate, gradually narrowed outwardly, the apex
truncate. Ninth sternite, 9s, very slender, expanded outwardly,
the apex deeply bilobed by a U-shaped notch, the slender lobes
with two long setae, one apical, the second placed more laterally
at base. Basistyle, b, with the apical spinous seta a little shorter
than its long basal tubercle. Dististyle, d, complex, the outer
arm at apex produced mesad at a right angle into a long black-
ened spike, with a smaller curved black spine at bend of outer
margin ; intermediate arm smaller but of somewhat similar shape
302 The Philippine Journal of Science
to the outer arm ; inner arm long, armed with groups of spines as
illustrated.
In colona (Plate 3, fig. 46) the apical lobe of the tergite, 9t, is
slightly longer, with the end gently emarginate. Ninth ster-
nite, 9s, with the lateral margins straight, the apex more gently
emarginate. Outer arm of dististyle, d, without a curved black
spine at angle; inner arm of very different conformation, as
shown.
Mindanao, Davao district, Calian, July 16, 1930 (C. F.
Clagg) ; holotype, male.
The distinctions between the present species and Styringomyia
colona Edwards (Krakatau) are best shown in the structure of
the male hypopygium.
ILLUSTRATIONS
[Legend: o, aedeagus ; 6, basistyle; d; dististyles ; db, dorsal lobe of basistyle; dd, dorsal dis-
tistyle; g, gonapophysis ; id, inner dististyle; od, outer dististyle; p, phallosome; e, 9th
Bternite; tt 9th tergite ; vb, ventral lobe of basistyle; vd; ventral dististyle.]
Plate 1
Fig. 1. Dolichopeza (Mitopeza) rizalensis sp. nov., wing.
2. Dolichopeza (Nesopeza) melanosterna sp. nov., wing.
3. Dolichopeza (Nesopeza) tarsalis Alexander, wing, medial field.
4. Dolichopeza (Mitopeza) mjobergi Edwards, wing, medial field.
5. Limonia (Limonia) bilobulifera sp. nov., wing.
6. Limonia (Limonia) melanopleura sp. nov., wing.
7. Limonia (Limonia) tremula sp. nov., wing.
8. Limonia (Libnotes) unistriolata sp. nov., wing.
9. Limonia (Libnotes) melancholica sp. nov., wing.
10. Limonia (Libnotes) perrara sp. nov., wing.
11. Limonia (Dicranomyia) orthia sp. nov., wing.
12. Limonia (Dicranomyia) neopunctulata sp. nov., wing.
13. Helius (Eurhamphidia) fuscofemoratus sp. nov., wing.
14. Helius (Eurhamphidia) indivisus sp. nov., wing.
15. Trentepohlia (Mongoma) distalis sp. nov., wing.
16. Gonomyia (Progonomyia) terebrella sp. nov., wing.
17. Erioptera (Teleneura) melanot&nia sp. nov., wing.
18. Erioptera (Empeda) lunensis sp. nov., wing.
19. Molophilus banahaoensis sp. nov., wing.
20. Molophilus procericomis sp. nov., wing.
21. Styringomyia armata Edwards, wing.
22. Styringomyia claggi sp. nov., wing.
Plate 2
Fig. 23. Dolichopeza (Mitopeza) rizalensis sp. nov., male hypopygium.
24. Dolichopeza (Nesopeza) melanosterna sp. nov., male hypopygium.
25. Limonia (Limonia) bilobulifera sp. nov., male hypopygium.
26. Limonia (Limonia) melanopleura sp. nov., male hypopygium.
27. Limonia (Libnotes) unistriolata sp. nov., male hypopygium.
28. Limonia (Libnotes) melancholica sp. nov., male hypopygium.
29. Limonia (Dicranomyia) orthia sp. nov., male hypopygium.
30. Limonia (Dicranomyia) neopunctulata sp. nov., male hypopygium.
31. Limonia (Dicranomyia) punctulata de Meijere, male hypopygium.
32. Limmonia (Dicranomyia) fullowayi Alexander, male hypopygium.
33. Helius (Eurhamphidia) indivisus sp. nov., male hypopygium.
34. Erioptera (Teleneura) melanotssnia sp. nov., male hypopygium.
262412 — io 303
304 The Philippine Journal of Science
Plate 3
Fig. 35. Erioptera (Empeda) lunensis sp. nov., male hypopygium.
36. Molophilus banahaoensis sp. nov., male hypopygium.
37. Molophilus procericomis sp. nov., male hypopygium.
38. Molophilus mendicus sp. nov., male hypopygium.
39. Molophilus tawagensis sp. nov., male hypopygium.
40. S tyring omyia luteipennis sp. nov., male hypopygium.
41. Styringomyia montina sp. nov., male hypopygium.
42. Styringomyia armata Edwards, male hypopygium.
43. Styringomyia claggi sp. nov., male hypopygium.
44. Styringomyia nigrostemata sp. nov., male hypopygium.
45. Styringomyia neocolona sp. nov., male hypopygium.
46. Styringomyia colona Edwards, male hypopygium.
Alexander: Philippine Tipulid^e, XL]
fPHTLlP. JOURN. Sci., 46, No. 2.
PLATE 1.
Alexander: Philippine Tipulid;e, XL]
[Philip. Journ. Sol, 46, No. 2.
PLATE 2.
Alexander: Philippine Tipulid^:, XL]
[Philip. Journ. Sci., 46, No. 2.
PLATE 3.
The Philippine
Journal of Science
Vol. 46 NOVEMBER, 1931 No. 3
AVIAN MALARIA STUDIES, I
PROPHYLACTIC PLASMOCHIN IN INOCULATED AVIAN MALARIA x
By Paul F. Russell
Of the International Health Division, Rockefeller Foundation
TWENTY TEXT FIGURES
INTRODUCTION
The drug plasmochin, sometimes spelled plasmoquine, was
developed in Germany in the Elberfeld laboratories of the I. G.
Farbenindustrie in 1926 by the chemists H6rlein,(65,66) Schule-
mann, Wingler, and Schonhofer(i58) working in close coopera-
tion with Roehl,(l49, 151) who used canaries as his experimental
animals. In the five years since then a great deal of attention
has been given this synthetic product in many laboratories
throughout the world. In the accompanying bibliography are
listed 194 plasmochin references and the list is not complete
as to Continental and South American periodicals. It is rather
remarkable that among these numerous publications there ap-
pear to be only three that refer to the possibility of using plas-
mochin to prevent malaria infection in man or birds.
1 In the examination of blood smears in the experiments reported in
parts I to IV of this series the author was assisted by Misses Amparo
Capistrano and Filomena Villacorta, microscopists on the staff of malaria
investigations of which the author is chief. This organization is sup-
ported by the Bureau of Science, Manila, in cooperation with the Interna-
tional Health Division of the Rockefeller Foundation. The experiments
were done at the Bureau of Science. This article was submitted for pub-
lication February 17, 1931.
263774 305
306 The Philippine Journal of Science 1931
Hegner and Manwell(60) by administering plasmochin to birds
in daily oral doses of 1.5 milligrams kept the blood of one bird
free from parasites for forty days after inoculation "with one
possible exception." Daily oral doses of 1.0 and 0.5 milligram
for five days after a single inoculation did not prevent the
appearance of parasites in the blood of birds.
Fischer (48) reported using plasmochin as a prophylactic drug
in man. He gave plasmochin to the crew of a ship calling at
West African river ports, and although he had 15 per cent
malaria morbidity, he contrasts this with 30 per cent on two
similar ships. Ejercito(44) in the Philippines gave prophylactic
plasmochin compound to eight individuals and prophylactic
quinine to eight others. Two of the first group and six of the
second acquired malaria during an eight weeks' test during
which they were not under strict control. He concluded that
plasmochin compound is apparently efficacious when used as a
prophylactic against malaria and maintains more subjects nega-
tive to malaria than quinine alone. He gave daily doses of 0.01
gram of plasmochin combined in tablet form with 0.125 gram
quinine sulphate to the first group and 10 grains of quinine
sulphate daily to the second.
A number of references may be found to a tendency that plas-
mochin seems to have to attack gametocytes in such a way that
they become devitalized and noninfective to mosquitoes.
Consult, for example, Roehl;(i49, 151) Green; (56) Manson-
Bahr;(97,98) Barber, Komp, and Newman ;(ii) and Whitmore,
Roberts, and Jantzen.(l93) There seems to be no doubt that
plasmochin has a genuine usefulness in malaria therapy, although
the tendency is to recommend that it be combined with quinine
for greater safety and effectiveness; consult, for example,
Green, (56) Manson-Bahr,(97,98) and Sinton.(l65)
The paucity and yet suggestiveness of the evidence as regards
prophylactic properties of plasmochin led to the experiments
reported in this paper.
GENERAL PROCEDURE
In the experiments here reported female canaries (Serinus
canarius) were used. These birds were purchased from local
dealers and were susceptible to the Plasmodium involved. Fe-
male birds were used because they cost less than males. The
parasite, Plasmodium cathemerium (Hartman, 1927), in over
two hundred cases has invariably established itself in these
46,3
Russell: Avian Malaria Studies, I
307
308 The Philippine Journal of Science 1931
birds upon injection, except during the periods of plasmochin
administration as noted below. This parasite was obtained
through the courtesy of Dr. C. G. Huff and is his "Boston
strain." It is not in direct line from the original isolation by
Hartman in 1924 from a Baltimore sparrow but was taken by
Huff from a Boston sparrow. The same strain of parasite was
used in all of the experiments reported in this and the second
paper of this series. The lines of transmission are shown in
fig. 1. All birds were kept well screened.
TECHNIC OF BLOOD INOCULATIONS
The technic of inoculation is simple. A vein on the inner
side of the left leg of the donor bird is punctured gently with a
Hagedorn needle, and blood is drawn into a 1-cubic-centimeter
tuberculin syringe half full of physiologic saline solution. After
each drop or two of blood is drawn into the syringe some of
the resulting mixture is ejected into a small vial that also
contains a little of the saline solution. By repeating this process
one soon has 1 or 2, or even 3, cubic centimeters, as required,
of a bright pink mixture of blood and saline solution, which is
thoroughly mixed. Using a 27-gauge needle, an injection (in
these experiments) of 0.3 cubic centimeter of the mixture was
made into the left breast muscle of the recipient bird. It is
possible also to infect birds by intraperitoneal and intravenous
routes, but in these experiments only intramuscular injections
were made.
USUAL COURSE OP INFECTION
After a prepatent period, which with P. cathemerium is usu-
ally from four to seven days, smears of the peripheral blood
as a rule show parasites in small numbers for three or four
days and then in great numbers for four or five days. If death
does not occur the blood stream then rapidly becomes relatively
free from parasites, but the blood usually remains infective
during the life of the bird, even over a period of years. Only
occasionally can parasites be demonstrated in smears; but in
this chronic or latent stage, even when parasites cannot be
demonstrated by prolonged microscopical examination, the blood
remains infective. Consult, for example, Wasielewski,(l89)
Sergent and Sergent, (160) and Whitmore;(i92) or take the case
of bird X36 in one of my experiments, typical of others in the
series. This bird became positive nine days after receiving
an injection of 0.3 cubic centimeter of blood-saline mixture
46,3 Russell: Avian Malaria Studies, I 309
taken as described above, from bird J53. A 30-minute examina-
tion of a blood smear from this donor bird (J53) taken the
day before, again at the time it was being bled for the inocula-
tion of X36, and the next day, showed no parasites at all.
SUPERINFECTION
If a bird be reinoculated with the same strain of Plasmo-
dium there is no superinfection. The bird is immune to a new
infection of any given strain so long as it has a chronic infec-
tion, and this, in most cases, means for the rest of its life. If,
however, the bird becomes entirely free of the Plasmodium in
question it can be reinfected. This phenomenon of immunity
to superinfection with the same species of Plasmodium is well
known to all who have studied avian malaria. Consult, for
example, Wasielewski,(i89) Moldovan,(H5) Sergent and Ser-
gent,(l60) and Taliaferro and Taliaferro. (175)
In these experiments this fact of immunity to superinfection
was used as a test to prove that the plasmochin-protected birds
were actually free from the Plasmodium injected into their
muscle. Had they been carrying parasites hidden from blood-
smear examination they would have been immune to subsequent
inoculations with the same strain of Plasmodium.
ADMINISTRATION OF PLASMOCHIN
The drug as used in these experiments was plasmochin sim-
plex, manufactured by I. G. Farbenindustrie A. G., Leverkusen,
Germany, for the Winthrop Chemical Company, Inc., New York.
It was purchased at a local pharmacy in boxes of ten ampoules
of 1 cubic centimeter each. According to the label the ampoules
contained a 1 per cent solution of plasmochin simplex, 2V-diethyl-
amino-isopentyl-8-amino-6-methoxy-quinoline. In other words
1 cubic centimeter of the solution contained 0.01 gram of plas-
mochin. To 1 cubic centimeter of this solution were added 4
cubic centimeters of distilled water. Thus, 5 cubic centimeters
of the resulting solution contained 0.01 gram of plasmochin.
In the first two experiments reported here, 0.1 cubic centi-
meter of this diluted solution was used as a daily dose; that is,
0.0002 gram of plasmochin simplex. This was given intramus-
cularly in the right breast, on the opposite side to that used for
the parasite inoculation.
310
The Philippine Journal of Science
1981
In the third experiment here recorded and in a fourth de-
scribed in the second paper of this series the dose of plasmochin
simplex was 0.00016 gram.
In some cases in the first two experiments there was necrosis
at the site of injection; but by making the injections well ante-
rior to avoid hsematomata and by inserting the needle deep in
the muscle and holding it steady during the injection, necrosis
was prevented in the last two experiments and the birds toler-
ated the injections very well.
In the first two experiments the mortality for a period of ten
days after the first injection of plasmochin among the twenty-
one birds used was 71 per cent. In the eleven control birds
(receiving no plasmochin) during the same period it was 45
per cent. In the last two experiments with improved technic,
a smaller dose of plasmochin, and probably a stronger lot of
birds, the mortality for ten days in the twenty birds receiving
plasmochin was 15 per cent and in the control group of fourteen
birds it was 29 per cent (see Tables 1 and 2) .
Table 1. — Mortality in first, second, and third experiments.
Num-
ber of
birds.
Died in 10
days or less.
Died in 15
days or less.
Died in 30
days or less.
Alive after
176 days or
more.
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
First experiment:
Plasmochin series
6
7
3
3
50
43
4
4
67
57
5
7
83
100
1
0
7
0
C on trols
Total
13
6
46
8
62
12
92
1
8
Second experiment:
Plasmochin series
15
4
12
2
80 i 13
87
75
14
3
93
75
(b)
Controls
50
3
Total
19
14
74
16
84
17
89
Third experiment:
Plasmochin series 8
Controls
10
4
2
0
20
0
2
0
20
0
4
2
40
50
1
2
10
50
Total
14
2
14
2
14
6
43
3
21
Totals for three experiments:
Plasmochin series
31
15
17
5
55
33
19
7
61
47
23
12
74
80
2
2
6
3
Controls __
Total _ __
46
22
48
26
57
35
76
4
9
a Six lived more than forty-five days.
b One lived thirty-four days.
c One lived seventy-seven days.
46,3 Russell: Avian Malaria Studies, I 311
Table 2. — Mortality, first and second versus third and fourth experiments.
N um-
ber of
Died in 10
days or less.
Died in 15
days or less.
Died in 30
days or less.
Alive after
170 days or
more.
experi-
birds.
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
81
15
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
Plasmoehin series:
First and second
ments
21
20
15
3
71
15
17
3
19
7
20
35
1
5
5
25
Third and fourth
ments.
experi-
Total.. _.
41
11
14
18
5
4
44
45
29
20
7
5
49
63
36
26
10
10
63
90
71
6
0
4
15
0
29
Controls:
First and second
ments
Third and fourth
ments,
experi-
experi-
Total
25
32
34
9
20
« 7
36
63
21
12
24
8
48
75
42
20
29
17
80
91
50
4
1
9
16
3
26
Totals:
First and second
ments
experi-
Third and fourth
ments
experi-
Total _.
66
27
41
32
48
46
70
10
15
The size of the dose was determined by the fact that 0.0002
gram was the largest amount of plasmoehin simplex which
would not cause signs of drug absorption in canaries. Increas-
ing the dosage caused signs beginning with unsteadiness of
gait and progressing as the dose became larger to coma and
death. (See the second paper of this series for a discussion of
the minimum lethal dose.)
BLOOD EXAMINATION
Blood smears were stained with Giemsa's stain and were
examined until a parasite was seen or, if none was seen, up
to a total of thirty minutes. If no parasites were found in
thirty minutes, the slide was called negative. If positive, it
was classified in accordance with the following scheme:
+ Positive in thirty minutes or less.
+ + Two parasites per field found more than twice in one min-
ute.
+ + + Three parasites per field found more than three times in
one minute.
^_ _^_ _|_ -_|. Four parasites per field found more than four times in one
minute.
H — I — | — | — I- Ten or more parasites per field on the average.
312
The Philippine Journal of Science
1931
This practical method of classification is suitable for this ex-
periment. The fact that it is a fairly good grouping may be
seen from Table 3, which also serves to present evidence as to
the approximate meaning of the plus signs.
Table 3. — Intensity grouping of blood smears.
Group.
+ .
+ + .
+ + + .
+ + ++.
+ + + + +
Number
of smears
counted.
74
19
18
15
44
Parasites
counted
per 10,000
red blood
cells.
15
170
320
560
1,320
FIRST EXPERIMENT (JUNE 19 TO JULY 21, 1930)
In the first experiment, as shown in fig. 2, plasmochin injec-
tions were started in six birds, U3, U12, U27, U28, U29, and
U33. Of these, only three, U12, U33, and U29, lived beyond the
first ten days of the experiment. There were seven controls,
U2, U25, U30, U31, U32, U34, and U35, of which two, U25 and
U34, died within ten days. The others all developed typical in-
fections. Of the three birds receiving plasmochin two, U12
and U29, died before they could be proved susceptible to malaria.
In the case of U12 blood was taken ten days after the attempt to
infect it. This blood proved to be noninfective to bird U40,
which twenty-seven days later was proved to be susceptible to
the same species of Plasmodium. In the case of U29 blood was
taken eight days after the attempt to infect it. This blood
proved to be noninfective to U42, which twenty-seven days later
was proved to be a susceptible bird.
There follow the protocols of U12, U33, and U29, which are
illustrated in figs. 3, 4, and 5.
FIRST EXPERIMENT
Protocol 1. Bird U12.
June 19, 1930. Blood smear negative from U12 (30 minutes).
June 28. Blood smear negative from U12 (30 minutes).
June 30 to July 6. U12 received 0.0002 gram plasmochin simplex by
intramuscular injection each day into right breast.
July 2. U12 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird 33R, known to be infective.
46,3
Russell: Avian Malaria Studies, I
313
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D = DIED
I s BLOOD INJECTED FROM
P s PLASM0CHIN SIMPLEX
t s PLASMODIA IN BLOOD SMEAR
0 s NO PLASMODIA FOUND IN BLOOD SMEAR
Fig. 2. Plasmochin simplex, a prophylactic drug: in avian malaria. First experiment.
An equal amount of the same mixture was given at the same time to
bird U30, as a control. Both injections made into left breast muscle.
Control bird, U30, became + July 10 and died July 23 of severe malaria.
July 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, and 21.
Daily blood smears from U12 negative. (Each smear searched for 30 min-
utes.)
July 12. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U12 injected into U40 which had
314
The Philippine Journal of Science
1933
JULY 2
— PLASMOCHIN SIMPLEX
JUNE 30-JULY 6
Fig. 3. Bird U12.
negative blood smears July 12, 18, 19, 20, 21,, 22, 23, 24, 25, 26, 28, 29, and
31, and August 1, 5, and 7. U40 became + August 8, which was seven days
after injection from bird 25R, which was known to be infective.
July 18. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U12 injected into U47, which died
July 23. (Of no value in this experiment.)
July 18. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from bird U32, which was known to
be infective, injected into ZJ12.
July 21. U12 died with no evidence of malaria.
Protocol 2. Bird U29.
June 30, 1930. Blood smear from U29 negative (30 minutes).
July 1. Blood smear from U29 negative (30 minutes).
June 30 to July 6. U29 received 0.0002 gram plasmochin simplex by in-
tramuscular injection each day into right breast.
July 4. U29 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird 33R, known to be infective.
An equal amount of the same mixture was given at the same time to bird
U32, as a control. Both injections were made into left-breast muscle.
The control bird, U32, became -f July 11 and died July 26 after severe
malaria.
July 7, 8, 9, 11, 12, 13, and 14. Daily blood smears from U29 negative.
(Each smear searched for 30 minutes.)
July 12. Three-tenths cubic centimeter physiologic saline solution mixed
with 5 to 7 drops of blood from U29 injected into U42, which had neg-
40,3
Russell: Avian Malaria Studies, I
315
Fig. 4. Bird U29.
ative blood smears July 12, 18, 19, 20, 21, 22, 23, 24, 25, 26, 28, 29, and
31, and August 1, 4, 5, and 7. U42 became + August 8, which was four
days after injection from bird U65, which was known to be positive.
July 14. Three-tenths cubic centimeter physiologic saline solution mixed
with 5 to 7 drops of blood from U29 injected into U43, which died July
15. (Of no value in this experiment.)
July 14. U29 died.
Physiologic saline mixture of bone marrow from U29 injected into U46,
whose blood smear was negative July 20 and Which died July 21.
Physiologic saline mixture of spleen pulp from U29 injected into U45,
which died July 17. (Of no value in this experiment.)
Protocol 3. Bird U33.
July 3, 1930. Blood smear from U33 negative (30 minutes).
July 3 to 8. TJ88 received 0.00016 gram plasmochin simplex by intra-
muscular injection about 10 a. m. each day into right breast.
July 5. U83 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird 33R, known to be infective.
An equal amount of the same mixture was given at the same time to bird
316
The Philippine Journal of Science
1931
« PLASMOCHIN SIMPLEX
JULY 3- 8
Fig. 5. Bird U33.
U35, as control. All injections were made into left breast muscle. U35
became positive July 12 and died July 14 of severe malaria.
July 8, 9, 10, 11, 12, 13, 14, 15, 16, 24, 28, 29, 30, and 31, and August 1,
2, 4, and 8. Daily blood smears from U33 negative. (Each smear searched
for 30 minutes.)
July 12. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U33 injected into U41, which had
negative blood smears July 12 to August 7. U41 became positive August 8,
which was four days after injection from U65, which was known to be
positive.
July 18. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U33 injected into U48, which had
negative blood smears July 19 to August 6. U48 became positive August
14, which was six days after injection from U42, which was known to be
positive.
August 4. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from U33 injected into U73, which
had negative blood smears July 31 to August 18. XJ73 became positive
46,3 Russell: Avian Malaria Studies, I 317
August 25, which was seven days after injection from U48, which was
known to be positive.
August 4. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird U65, known to be infective,
injected into U38.
August 9, U33 +; August 11, U83 + + ; August 12 and 13, U33 + + + ;
August 14 to 28, U33 +; October 11, U33 +.
SECOND EXPERIMENT (JULY 19 TO AUGUST 24, 1930)
Encouraged by the one clear-cut success in the first experi-
ment, a series of fifteen birds, U50 to U64, were given plas-
mochin simplex in 0.0002 gram intramuscular doses, as described
above, every day for seven days. Infected blood was injected
into these birds and into four control birds, U7, U65, U66, and
U67, on the third day. Two of the controls, U7 and U66, and
ten of the birds receiving plasmochin, U50 to U52, U57 to U59,
and U62 to U64, died within ten days and were of no value
in the experiment. The other two controls had typical malaria
infections (see fig. 6).
Of the five remaining birds receiving plasmochin two, U56
and U60, lived thirty-five and thirty-one days, respectively; long
enough to give clear-cut results. The other three died before
they could be proved to be susceptible. None showed any evi-
dence whatever of malaria after thirteen, thirteen, and fourteen
days, respectively. In the case of U53 and U61 blood was taken
on the ninth day after attempted infection. This blood proved
to be noninfective when injected into birds that in each case
were proved to be susceptible twenty-eight days later.
The protocols of U53, U54, U56, U60, and U61 follow. These
protocols are illustrated in figs. 6 to 10, inclusive.
SECOND EXPERIMENT
Protocol 1. Bird U53.
July 19, 1930. Blood smear from U53 negative (30 minutes).
July 19 to 24. JJ53 received 0.0002 gram plasmochin simplex by in-
tramuscular injection each day about 10 a. m. into right breast.
July 21. U58 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from U32, known to be infective. An
equal amount of the same mixture was given at the same time to birds
U64, U65, U66, and U67, as controls. All injections were made into left
breast muscle about 3 p. m. Control bird U64 died, negative, July 26;
U65 became + July 27 and died August 5 of severe malaria; U66 died,
negative, July 27; U67 became + July 27 and had a mild infection.
July 28, 29, 30, and 31. Daily blood smears from USS negative. (Each
smear searched for 30 minutes.)
318
The Philippine Journal of Science
1931
Fig. 6. Plasmochin simplex, a prophylactic drug in avian malaria. Second experiment.
July 30. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U53 injected into U68, which had
negative blood smears July 31, and August 7, 8, 9, 11, 12, 14, 15, 16, 18, and
20. U68 became + August 27, which was six days after injection from
bird U89, which was known to be infective.
July 31. U5S died with no evidence of malaria.
46, ;
Russell: Avian Malaria Studies, I
319
Fig. 7. Bird U53.
Protocol 2, Bird U54.
July 19, 1930. Blood smear from TJ5U negative (30 minutes).
July 19 to 24. TJ5Jt. received 0.0002 gram plasmochin simplex by intra-
muscular injection each day about 10 a. m. into right breast.
July 21. U54. received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U32, known to be infective.
An equal amount of the same mixture was given at the same time to birds
U64, U65, U66, and U67, as controls. All injections were made into left
breast muscle about 3 p. m. Control bird U64 died, negative, July 26;
U65 became + July 27 and died August 5 of severe malaria; U66 died,
negative, July 27; U67 became + July 27 and had a mild infection.
July 28, 29, 30, and 31. Daily blood smears from U5U negative. (Each
smear searched for 30 minutes.)
July 30. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U54 injected into U70, which was neg-
ative August 7, 8, and 9 and died August 10 with no evidence of malaria.
July 31. U5Jt died with no evidence of malaria.
320
The Philippine Journal of Science
1931
Fig. 8. Bird U54.
Protocol 3. Bird U56.
July 19, 1930. Blood smear from U56 negative (30 minutes).
July 19 to 24. U56 received 0.0002 gram plasmochin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
July 21. U56 received 0,3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U32, known to be infective.
An equal amount of the same mixture was given at the same time to
birds U64, U65, U66, and U67, as controls. All injections were made into
left breast muscle. U65 and U67 became positive July 27. U64 and U66
died July 26 and 27, respectively.
July 28, 29, 30, and 31, and August 1, 2, 4, 5, 6, 7, 8, 9, 11, 14, and 15.
Daily blood smears from U56 negative. (Each smear searched for 30
minutes.)
July 30. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U56 injected into U72, which had
negative blood smears July 31 to August 20. U72 became positive August
27, which was six days after injection from U89, which was known to be
positive.
46,3
Russell: Avian Malaria Studies, I
321
Fig. 9. Bird U56.
August 12. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U56 injected into U92, which had
negative blood smears August 12 to 26. U92 became positive September
1, which was six days after injection from U91, which was known to be
positive.
August 12. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird U40, known to be infective,
injected into TJ56.
August 16 and 18, U56 +; August 19, U56 + + + ; August 20, U56 + + ;
August 21, U56 + + + ; August 22, U56 ++■'; August 23, U56 +; August
24, died.
Protocol 4. Bird U60.
July 19, 1930. Blood smear from U60 negative (30 minutes).
July 19 to 24. U60 received 0.00016 gram plasmochin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
July 21. U60 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U32, known to be infective.
An equal amount of the same mixture was given at the same time to birds
263774 2
322
The Philippine Journal of Science
1931
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Fig. 10. Bird U60.
U64, U65, U66, and U67, as controls. All injections were made into
left breast muscle. U65 and U67 became positive July 27. U64 and U66
died July 26 and 27, respectively.
July 28, 29, 30, and 31, and August 1, 2, 4, 5, 6, 7, 8, 9, 11, 14, and 15.
Daily smears from U60 negative. (Each smear searched for 30 minutes.)
July 30. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U60 injected into U71, which had
negative blood smears August 7 and 8 and died August 8.
August 5. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U60 injected into U22, which had
negative blood smears June 19 to August 19. U22 became positive August
26, which was six days after injection from U88, which was known to be
positive.
August 12. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from U60 injected into U93, which
had negative blood smears August 12 to 26. U93 became positive Sep-
tember 1, which was six days after injection from, U91, which was known
to be positive.
(August 12. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird U40, known to be infective,
injected into U60.
46,3
Russell: Avian Malaria Studies, I
323
August 16 and 18, U60 -f ; August 19 and 20, U60 +-f »+ ; August 20,
U60 died.
Protocol 5. Bird U61.
July 19, 1930. Blood smear from U61 negative (30 minutes).
July 19 to 24. U61 received 0.0002 gram plasmochin simplex by intra-
muscular injection each day about 10 a. m, into right breast.
July 21. U61 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U32, known to be infective.
An equal amount of the same mixture was given at the same time to
birds U64, U65, U66, and U67, as controls. All injections were made
into left breast muscle about 3 p. m. Control bird U64 died, negative, July
26; U65 became -f July 27 and died August 5 of severe malaria; U66
died, negative, July 27; U67 became + July 27 and had a mild infection.
July 28, 29, 30, and 31, and August 1. Daily blood smears from U61
negative. (Each smear searched for 30 minutes).
July 30. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from U61 injected into U69, which had
negative blood smears July 31, and August 7, 8, 9, 11, 12, 14, 15, 16, 18,
and 20. U69 became + August 27, which was six days after injection
from U89, which was known to be infective.
August 2. U61 died with no evidence of malaria.
Fig. 11. Bird U61.
324 The Philippine Journal of Science 1931
THIRD EXPERIMENT (AUGUST 9 TO OCTOBER 9, 1930)
Using a smaller dose, 0.00016 gram, as explained above, ten
canaries, U78 to U87, were given daily intramuscular injec-
tions of plasmochin and were injected with infected blood on
the third day (see fig. 11). Four controls, U88 to U91, were
also injected with the same amount of the same blood taken
at the same time. All of the controls developed typical malaria.
Two of the birds receiving plasmochin, U82 and U83, died within
ten days and were of no use in the experiment. The other
eight all lived long enough to demonstrate clearly that their
plasmochin injections had prevented malaria. Each of the four
birds U78, U79, U80, and U84, after being negative for twenty-
one days, was proved to be susceptible to the Plasmodium used
in the first injection. Four birds, U81, U85, U86, and U87,
remained negative for forty-four days each and were then
proved to be susceptible. Blood taken from U78, U79, U81,
U85, and U87 on the twenty-first, sixteenth, fourteenth, and
forty-fourth days, respectively, proved to be noninfective to
birds that in each case after two weeks were proved to be sus-
ceptible birds.
There follow the protocols of birds U78, U79, U80, U81, U84,
U85, U86, and U87. These are illustrated by figs. 12 to 19,
inclusive.
THIRD EXPERIMENT
Protocol 1. Bird U78.
August 9, 1930. Blood smear from U78 negative (30 minutes).
August 9 to 15. U78 received 0.00016 gram plasmochin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
August 11. U78 received 0.3 cubic centimeter physiologic saline so-
lution containing 5 to 7 drops of blood from bird U42, known to be in-
fective. An equal amount of the same mixture was given at the same
time to birds U88, U89, U90, and U91, as controls. All injections were
made into left breast muscle. U88 and U89 became positive August 16;
U90 and U91, August 18. U91 died August 28 of severe malaria.
August 18, 19, 21, 23, 25, 27, and 29. Daily smears from U78 nega-
tive. (Each smear searched for 30 minutes.)
September 1. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from U78 injected into U97, which
had negative blood smears September 1, 6, 9, 11, and 13. U97 became +
September 22, which was seven days after injection from U80, which was
known to be positive.
September 1. Three-tenths cubic centimeter physiologic saline solu-
tion mixed with 5 to 7 drops of blood from bird U69, known to be in-
fective, injected into ZJ78.
September 6 and 8, U78 + ; September 9, 10, 11, 12, and 13, U78
-I- + -|> _|_ 4. j September 14, U78 died of acute malaria.
46,3
Russell: Avian Malaria Studies, I
325
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Fig. 12. Plasmochin simplex, a prophylactic drug in avian malaria. Third experiment.
Protocol 2. Bird U79.
August 9, 1930. Blood smear from U79 negative (30 minutes).
August 9 to 15. U79 received 0.00016 gram plasmochin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 11. U79 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U42, known to be infective.
An equal amount of the same mixture was given at the same time to birds
U88, U89, U90, and U91, as controls. All injections were made into left
326
The Philippine Journal of Science
1931
Fig. 13. U78.
breast muscle. U88 and U89 became positive August 16. U90 and U91
became positive August 18. U91 died August 28 of severe malaria.
August 19, 21, 23, 25, 27, and 29, and September 1. Daily blood
smears from U79 negative, (Each smear searched for 30 minutes.)
September 1. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from U79 injected into U98, which
had negative blood smears September 1 to 15. U98 became positive Sep-
tember 22, which was seven days after injection from U80, which was
known to be positive.
September 1. Three-tenths cubic centimeter physiologic saline solu-
tion mixed with 5 to 7 drops of blood from bird U69, known to be infec-
tive, injected into U79.
September 6 to 8, U79 + ; September 9, U79 +\+; September 10, U7&
+:+!+ + + ; September 11, U79 + + ; September 12 to 16, U79 -f ; Octo-
ber 3, U79 died.
46,3
Russell: Avian Malaria Studies, I
327
FIG. 14. Bird U79.
Protocol 3. Bird U80.
August 9, 1930. Blood smear from U80 negative (30 minutes).
August 9 to 15. U80 received 0.00016 gram plasmochin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 11. U80 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U42, known to be in-
fective. An equal amount of the same mixture was given at the same
time to birds U88, U89, U90, and U91, as controls. All injections were
made into left breast muscle. U88 and U89 became positive August 16.
U90 and U91 became positive August 18. U91 died August 28 of severe
malaria.
August 18, 19, 21, 23, 25, 27, and 29, and September 1. Daily blood
smears from U80 negative. (Each smear searched for 30 minutes.)
September 1. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird U69, known to be infective,
injected into TJ80,
328
The Philippine Journal of Science
1931
Fig. 15. Bird TJ80.
September 6, U80 +; September 8, U80 +;+ + ; September 9, ZJ80
+ + ; September 10 and 11, U80 +; September 12, U80 + + ; September
13 and 15, U80 ++'+; September 16, U80 ++; September 17 to 19, ZJ80
+ ; October 3, U80 +.+ +!+ + ; October 5, U80, died.
Protocol 4. Bird U81.
August 9, 1930. Blood smear from JJ81 negative (30 minutes).
August 9 to 15. U81 received 0.00016 gram plasmochin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 11. U81 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U42, known to be in-
fective. An equal amount of the same mixture was given at the same
46,3
Russell: Avian Malaria Studies, I
329
time to birds U88, U89, U90, and U91, as controls. All injections were
made into left breast muscle. U88 and U89 became positive August 16.
U90 and U91 became positive August 18. U91 died August 28 of se-
vere malaria.
August 18, 19, 21, 23, 25, 27, and 29, and September 6, 8, 10, 12, 15, 17,
19, 22, and 24. Daily blood smears from U81 negative. (Each smear
searched for 30 minutes.)
September 6. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from U81 injected into 54R, which
had negative blood smears September 6 to 26. 54R became positive Sep-
tember 27, which was seven days after injection from J 10, which was
known to be positive.
September 24. Three-tenths cubic centimeter physiologic saline solu-
tion mixed with 5 to 7 drops of blood from bird U97, known to be in-
fective, injected into U81.
September 29 and 30, U81 +; October 1, U81 +; October 2, U81 ■+• + + ;
October 3 to 6, U81 ++'+ + + ; October 10 and 21, U81 +.
Fig. 16. Bird U81.
330
The Philippine Journal of Science
1931
Protocol 5. Bird U84.
August 9, 1930. Blood smear from U8k negative (30 minutes).
August 9 to 15. U81+ received 0.00016 gram plasmoehin simplex by
intramuscular injection about 10 a. m, each day into right breast.
August 11. U8U received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U42, known to be in-
fective. An equal amount of the same mixture was given at the same
time to birds U88, U89, U90, and U91, as controls. All injections were
made into left breast muscle. U88 and U89 became positive August 16.
U90 and U91 became positive August 18. U91 died August 28 of se-
vere malaria.
August 18, 19, 21, 23, 25, 27, and 29, and September 1. Daily blood
smears from TJ8U negative. (Each smear searched for 30 minutes.)
Fig. 17. Bird U84.
46,3 Russell: Avian Malaria Studies, I 331
September 1. Three-tenths cubic centimeter physiologic saline solu-
tion mixed with 5 to 7 drops of blood from bird U69, known to be in-
fective, injected into U84.
September 4 and 5, U84 + ; September 6, U8U -f -f -f ; September 8 to
11, U84 + + + + + ; September 11, U8J> died.
Protocol 6. Bird U85.
August 9, 1930, Blood smear from U85 negative (30 minutes).
August 9 to 15. U85 received 0.00016 gram plasmochin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 11. U85 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 dlrops of blood from bird U42, known to be in-
fective. An equal amount of the same mixture was given at the same
time to birds U88, U89, U90, and U91, as controls. All injections were
made into left breast muscle. U88 and U89 became positive August 16.
U90 and U91 became positive August 18. U91 died August 28 of se-
vere malaria.
August 18, 19, 21, 23, 25, 27, and 29, and September 1, 3, 5, 8, 10, 12,
15, 17, 19, 22, and 24. Daily blood smears from U85 negative. (Each
smear searched for 30 minutes.)
September 24. Three-tenths cubic centimeter of physiologic saline so-
lution mixed with 5 to 7 drops of blood from U85 injected into U99,
which had negative blood smears September 24 to October 19. U99 became
positive October 14, which was five days after injection from J42, which
was known to be positive.
September 24. Three-tenths cubic centimeter physiologic saline solu-
tion mixed with 5 to 7 drops of blood from bird U97, known to be in-
fective, injected into U85.
September 29 and 30 and October 1, U85 +; October 2, JJ85 + + ;
October 3 and 4, U85 +.+ + ; October 6, U85 -f ,+ ; October 10, U85 +;
October 18, U85 +.+ + + + ; October 18, U85 died.
Protocol 7. Bird U86.
August 9, 1930. Blood smear from U86 negative (30 minutes).
August 9 to 15. U86 received 0.00016 gram plasmochin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
August 11. U86 received 0.3 cubic centimeter physiologic saline so-
lution containing 5 to 7 drops of blood from bird U42, known to be in-
fective. An equal amount of the same mixture was given at the same
time to birds U88, U89, U90, and U91, as controls. All injections Were
made into left breast muscle. U88 and U89 became positive August 16.
U90 and U91 became positive August 18. U91 died August 28 of severe
malaria.
August 18, 19, 21, 23, 25, 27, and 29, and September 1, 3, 5, 8, 10,
12, 15, 17, 19, 22, 24, and 29. Daily blood smears from U86 negative.
(Each smear searched for 30 minutes.)
September 24. Three-tenths cubic centimeter physiologic saline solu-
tion mixed with 5 to 7 drops of blood from, bird U97, known to be infec-
tive, injected into XJ86.
332
The Philippine Journal of Science
1931
Fig. 18. Bird U85.
September 30 and October 1 and 2, U86 + ; October 3, U86 +1+ ; Oc-
tober 4, U86 ',+ + + +!+; October 6, U86 ++!+.; October 10, U86 +;
October 13, TJ86 died.
Protocol 8. Bird U87.
August 9, 1930. Blood smear from ZJ87 negative (30 minutes).
August 9 to 15. TJ87 received 0.00016 gram plasmoehin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 11. U87 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U42, known to be infective.
An equal amount of the same mixture was given at the same time to
birds U88, U89, U90, and U91, as controls. All injections were made into
left breast muscle. U88 and1 U89 became positive August 16. U90 and
U91 became positive August 18. U91 died August 28 of severe malaria.
46,8
Russell: Avian Malaria Studies, I
333
Fig. 19. Bird U86.
August 18, 19, 21, 23, 25, 27, and 29, and September 1, 3, 5, 8 10,
12, 15, 17, 19, 22, and 24. Daily blood smears from U87 negative. (Each
smear searched for 30 minutes.)
September 24. Three-tenths cubic centimeter of physiologic saline so-
lution mixed with 5 to 7 drops of blood from U87 injected into U100,
which had negative blood smears September 24 to October 9. U100 be-
came positive October 14, which was five days after injection from J42^
which was known to be positive.
September 24. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird U97, known to be infective,
injected into TJ87.
September 29 to October 1, TJ87 + ; October 2, US7 -f + ; October 3, U87
+ ■+ + + ; October 4 and 6, U87 + + + + + ; October 6, U87 died.
334
The Philippine Journal of Science
1931
Fig. 20. Bird U87.
SUMMARY
Three experiments are reported in which canaries were given
intramuscular injections of a mixture of saline and blood con-
taining Plasmodium cathemerium on the third day of a week
during which they received daily doses of plasmochin simplex
intramuscularly. In no case was it possible to detect an infec-
tion in these birds, although in every case control birds that
had not received plasmochin developed typical avian malaria.
CONCLUSION
It is concluded that the infection of a canary by experimental
needle inoculation with Plasmodium cathemerium (Hartman,
46'3 Russell: Avian Malaria Studies, I 335
1927) can be prevented by intramuscular injections of plasmo-
chin simplex in daily doses of from 0.00016 to 0.0002 gram.
author's note
These experiments were reported by the author in Bangkok
in December, 1930, as noted on page 32, paragraph 77, of the
8th Congress— Far Eastern Association of Tropical Medicine —
Abstracts of Papers and Programme of Scientific Sessions —
Bangkok, December 9 to 12, 1930.
Because of the important implications of these experiments
as regards human malaria they were also discussed and sum-
marized in a paper published by the American Journal of Trop-
ical Medicine in July, 1931.
That this emphasis was justified has been shown by the fact
that, on June 6, 1931, it was reported in the London Lancet,
volume 220, No. 5623, that James had protected not only birds
but also humans against malaria by using beprochin, a drug
probably identical with plasmochin. It would appear that once
again an experiment in avian malaria has been a reliable in-
dicator as regards human malaria.
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plasmochin. Arb. u. Tropenkrankli (Festschr. B. Nocht) (1927) 507-
511. (Note: Doctor Schulemann has edited a booklet called "Plas-
moquine, Plasmoquine Compound, Quino-plasmoquine," published in
1930 by the I. G. Farbenindustrie A. G. of Leverkusen, Germany.
This contains a large number of references.)
159. Segal, M., and J. Block. Plasmochin. Arch. f. Sehiffs- u. Trop.-
Hyg. 33 (1929) 532-535.
342 The Philippine Journal of Science mi
160. Sergent, Et and Ed. Ann. Inst. Past. 35 (1921) 125-141.
161. Serio, F. Plasmochin. Riv. di Malariol. 8 (1929) 436-448.
162. Sholle, G. G. Plasmoquinin (plasmochin) in children. Mosk. Med.
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163. Sinton, J. A. Treatment of malarial fevers. Trans. 7th Congr. Far
Eastern Assoc. Trop. Med. 2 (1927) 804-813.
164. Sinton, J. A., and W. Bird. Plasmoquine. Ind. Journ. Med. Res.
16 (1928) 159-177.
165. Sinton, J. A., S. Smith, and D. Pottenger. Plasmoquine and qui-
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166. Sioli, F. Effect of plasmochin in malarial superinfection. Natur-
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167. Sioli, F. Plasmochin in treatment of general paralysis. Beih. z.
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176. Tanew, I., and G. Haschnow. Value of plasmochin. Muench. Med.
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46,3 Rttssell: Avian Malaria Studies, I 343
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(1928).
ILLUSTRATIONS
TEXT FIGURES
Fig. 1. Diagram showing lines of transmission and attempted transmis-
sion of Plasmodium cathemerium. Experiments 1 to 4.
2. Plasmochin simplex, a prophylactic drug in avian malaria. First
experiment.
3. Bird U12.
4. Bird U29.
5. Bird U33.
6. Plasmochin simplex, a prophylactic drug in avian malaria. Second
experiment.
7. Bird U53.
8. Bird U54.
9. Bird U56.
10. Bird U60
11. Bird U61.
12. Plasmochin simplex, a prophylactic drug in avian malaria. Third
experiment.
13. Bird U78.
14. Bird U79.
15. Bird U80.
16. Bird U81.
17. Bird U84.
18. Bird U85.
19. Bird U86.
20. Bird U87.
345
AVIAN MALARIA STUDIES, II
PROPHYLACTIC PLASMOCHIN VERSUS PROPHYLACTIC QUININE
IN INOCULATED AVIAN MALARIA 1
By Paul P. Russell
Of the International Health Division, Rockefeller Foundation
SEVEN TEXT FIGURES
INTRODUCTION
In the first paper of this series (i) three experiments were
reported in which experimental inoculation of canaries with
Plasmodium cathemerium (Hartman, 1927) was invariably pre-
vented by intramuscular injections of plasmochin simplex in
daily doses of from 0.00016 to 0.0002 gram. In all cases the
attempt to infect the birds was made on the third day of the
series of plasmochin injections.
In the above-mentioned first paper a discussion was given of
plasmochin and an extensive bibliography was prepared. There
was also a full description of the technic of injection, of the
examination of blood smears, and of other pertinent phases of
the work.
The fourth experiment herein reported was along the same
general lines but with the following two notable changes in
procedure.
In the first place instead of attempting infection always on
the third day, in this case inoculations were made on various
days as noted below. Secondly, a parallel series of birds was
studied in which prevention was attempted by using quinine
instead of plasmochin.
In all other respects the technic followed was that described
in the first paper, (l) The mortality among the birds used in
this experiment is shown in Table 1.
1 Misses Amparo Capistrano and Filomena Villacorta, microscopists
on the staff of malaria investigations, assisted in the examination of blood
smears in this experiment. The work was done at the Bureau of Science,
Manila, with the cooperation of the International Health Division of the
Rockefeller Foundation.
347
348 The Philippine Journal of Science
Table 1. — Mortality of birds in fourth experiment.
1931
Num-
ber of
birds.
Died in 10
days or less.
Died in 15
days or less.
Died in 30
days or less.
Alive after
170 days.
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
Num-
ber.
Per
cent.
Plasmochin series
10
20
10
1
1)
4
10
5
40
1
6
5
10
30
50
3
11
8
30
55
80
4
6
2
40
30
20
30
Quinine series
Controls
Total _
40
6
15
12
30
22
55
12
FOURTH EXPERIMENT — PLASMOCHIN SERIES
(AUGUST 25 TO OCTOBER 20, 1930)
Ten canaries, Jl to J10, were each given an intramuscular
injection of plasmochin simplex, 0.00016 gram, each morning
for seven days at about 10 a. m. into the right breast muscle.
Two birds, Jl and J2, were inoculated with infected blood into
the left breast muscle at 3 p. m. of the third day. In a similar
way J2 and J4 were inoculated on the fourth day; J5 and J6
on the fifth day; J7 and J8 on the sixth day; J9 and J10 on
the seventh day (see fig. 1). The last pair, J9 and J10, there-
fore, received their inoculation of infected blood about five hours
after the last injection of plasmochin. These two birds devel-
oped malaria on the tenth and eleventh days after inoculation
and ran typical courses. The other eight birds, Jl to J8, all
remained negative. Two control birds were injected each day
with the same infected blood, in the same amount, and at the
same time as the birds that had received plasmochin. These
birds, J31 to J40, all became positive and had typical malaria,
with the exception of J35 and J37, which died on the third and
second days, respectively, after infection. (There remained at
least one control bird for each day of infection.)
Of the birds receiving plasmochin, J4 and J8 died on the
fifth and twelfth days, respectively, both negative. Jl, J2, J3,
J5, J6, and 31 remained negative for 26, 42, 25, 24, 40, and 39
days, respectively. Each was then proved to be a susceptible
bird by an injection of positive blood. Each had a typical
malaria course, J3 dying in the acute phase. Blood was taken
from some of the plasmochin birds at intervals to test its inf ec-
tiveness and invariably was noninfective, although in each case
the recipient was subsequently proved to be susceptible. Table
2 lists these tests of inf ectivity.
Saug
J.
1
2
3
4
5
6
7
8
9
10
II
12
13
14
15
16
17
18
19
20
21
22
23 24
25 26
27
28
29
3o|ai
I?
Up p
up F
P
_E
P
J3
m
P
P
^
P
p
Op p
p
J4
lUQj
"©p p
p
Op p
P
P
^
J5
__
N/7
"*P
P
p
P
up
P
P
J6
~
^
P
p
P
^
P
P
J8
__
^P
P
p
F
P
F
P
J9
-
^P
P
P
P
p
P
m
JIO
_
luii
~°P
p
F
' P
p
P
J3I
~
—
__
rj?
0
J 32
__
U9C
~o~
J33
—
~
-
-
-
-
-
LUdJ
h— -
0
jj34
-
IU9I
0
J 35
_
~
^
-
-
fU7j
0
-t*-
■~
^
_
__
_.
--
-
—
W73.
0
t J 37
"-
—
-
_.
— ■
SB7
0
| J 38
—
^
-
—
IU3'
0
—
._
._.
"-
-_
^
X>2'<
(
0
1 J 40
-
"
_
—
-
V21
— |
0
!: U 22 !
-
*H4-
, U37
-
"
-
-
_
—
U73
--
-
___
U90
-
._
_
~
-
—
^
_
-
U9I
_
—
--
-
_.
-
~-
__.
—J
SEPT
Jl
0-
<h
~
■-©-
._
p
-
luao
0
-
<y
J2
-
n
n
-Q--
"1
■
-o-
-o-
"*
"~*^
J3
-0
_
O
o~
<±-
!U8f
__
--+-
~+-
-H~
_.
— i
0
0
o
0
^
-0
J5
Q
P
p
n
U3t
0~
J6
p
n
0
p
p
0
p
P
p
_A_
rt
J7
Q
P
-
n
p
P
n
-O—
o-
"-
-o
_.
~
"
"
^tJ—
""
J8
-$-
-
--
__
__
1
°D
J3
o
0
--
-^
-
__
_
'p'
JIO
__
p
0
—
-+
■-+-
-
"
°
J3I
+
-»+
"
-"
-
—
_
T
J32
-+-
~$
—
"
-
-
-
1 i^a
JJi
^
++-
H~»+
+
_.
p
—
--
J34
6
|j J 36
-
.
1
"d"
! J38
-
—
1
mi
1
'
j;j39
0-
-
_.
-
|J41
__
._
r.ii
_
_ .
0
0
0
H>
0
!lj42
-
IJ2
0
_
-0
0
0
0
J43
--
IJ3
—
o
-
-
&
9
J44
LJ5
0
__
-
—
._
-
-
a
^
J45
__
._.
0
'
^
0
U80
J2
._
—
-
IM
,
-
D-
-
--
0
0
'
"
J6
154
?
—
0
+
*
+4-
+*
*♦♦
+4
-
0
J7
t54
?
0
J4I
IW
?
£^
--
0
*
'
1
J42
[54
,
n
(
!
llll
->--
P
_
-
__
---
—
_
O"
H
Q
J43
l«S4
^>
_
0
0
-tf-
hi i
1
J44
l«>4
?
-o-
-
0
— *
'
1
1
'
J45
54
*
0
J46
[J6
0
Ml
H+-
H»*-f
lit
A
+
+
n
o„
J4-7
,12
J43
«-
*
0
0,
J48
1,17
-
J4."
0
°
0
'
64R
V
2
3
4
5
6
7
8
9
10
H
»2
13
14
\b
16
17
is
19
20
2»
22
23
24
25
26
21
28
29
ao
31
Dr DIED
1 = BLOOD INJECTED FROM
P= PLASMOCHIN SIMPtEX
+ ^ PLASMODIA FOUND IN BLOOD SMEAR
0 = NO PLASMODIA FOUND IN BLOOD SMEAR
Fig. 1. Plasmochin simplex, a prophy
263774 Facing page 348.
actic drug in avian malaria. Fourth experiment.
46,3
Russell: Avian Malaria Studies, II
Table 2. — Tests of infectivity.
349
Donor.
Jl
J2
J2
J3
J5
J6
J6
J7
Days after
attempted
infection
of donor.
Recipient.
21
J41
21
J42
38
J47
21
J43
21
J44
21
J45
36
J46
35
J48
Result.
Negative.
Do.
Do.
Do.
Do.
Do.
Do.
Do.
There follow the protocols of birds Jl, J2, J3, J5, J6, J7, J8,
J9, and J10. Text figs. 2 to 7 illustrate the protocols of birds
Jl, J2, J3, J5, J6, and J7.
FOURTH EXPERIMENT
Protocol 1. Bird Jl.
August 23, 1930. Blood smear from Jl negative (30 minutes).
August 23 to 29. Jl received 0.00016 gram plasmochin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 25. Jl received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U90, known to be infective.
An equal amount of the same mixture was given at the same time to birds
J31 and J32, as controls. All injections were made into left breast muscle.
J31 and J32 became positive September 1. J31 died September 8 of severe
malaria.
August 25, and September 1, 3, 5, 8, 10, 12, 15, 18, and 20. Daily blood
smears from Jl negative. (Each smear searched for 30 minutes.)
September 15. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from Jl injected into J41, which
had negative blood smears September 15 to 26 and October 2 to 6. J41
became positive October 7, which was five days after injection from 54R,
which was known to be positive.
September 15. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird U80, known to be infective,
injected into Jl.
September 22 and 23, Jl + ; September 25, Jl -f -f- -f- -f ; September
26, Jl -j--f + + + ; September 29 and October 3, Jl +; October 20, Jl 0.
Protocol 2. Bird J2.
August 23, 1930. Blood smear from J2 negative (30 minutes).
August 23 to 29. J2 received 0.00016 gram plasmochin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
August 25. J2 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U90, known to be infective.
An equal amount of the same mixture was given at the same time to birds
J31 and J32, as controls. All injections were made into left breast muscle.
350
The Philippine Journal of Science
1931
Fig. 2. Bird Jl.
J31 and J32 became positive September 1. J31 died September 8 of severe
malaria.
August 25, September 2, 4, 6, 9, 11, 13, 15, 18, 20, 22, 24, 26, 29, and
October 2 and 6. Daily blood smears from J 2 negative. (Each smear
searched for 30 minutes.)
September 15. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from J2 injected into J42, which had
negative blood smears September 15 to October 4. J42 became positive
October 6, which was four days after injection from 54R, which was
known to be positive.
October 2. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from J2 injected into J47, which had
negative blood smears October 2 to 23. J47 became positive October 24,
which was eight days after injection from J43, which was known to be
positive.
46,3
Russell: Avian Malaria Studies, II
351
Fig. 3. Bird J2.
October 2. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird 54R, known to be infective, in-
jected into J2.
October 7 to 11, 32 +; October 13 and 14, 32 + + + + + ; October 15
and 16, J2 + +;+ + ; October 17, J 2 + + -|~f- + ; October 21, J2 died.
Protocol 3. Bird J3.
August 23, 1930. Blood smear from 33 negative (30 minutes).
August 23 to 29. 33 received 0.00016 gram plasmochin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 26. 33 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U91, known to be infective.
An equal amount of the same mixture was given at the same time to birds
J33 and J34, as controls. All injections were made into left breast muscle.
352
The Philippine Journal of Science
1931
J33 and J34 became positive September 1. J33 died September 8 of severe
malaria.
August 26, and September 1, 3, 5, 8, 10, 12, 15, 16, 18, and 20. Daily
blood smears from JS negative. (Each smear searched for 30 minutes.)
September 16. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from JS injected into J43, which
had negative blood smears September 16 to October 8. J43 became posi-
tive October 9, which was seven days after injection from 54R, which
was known to be positive.
September 16. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird U80, known to be infective,
injected into JS.
September 22, 23, J 3 +; September 25, J 3 -f-4-; September 26, JS
_l_ 4. _j_ _|_ _|_ ; September 29, JS died.
Fig. 4. Bird J3. Fourth experiment.
40,3
Russell: Avian Malaria Studies, II
353
Protocol 4. Bird J5.
August 23, 1930. Blood smear from J 5 negative (30 minutes).
August 23 to 29. J5 received 0.00016 gram plasmoehin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
August 27. J 5 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U73, known to be infective.
An equal amount of the same mixture was given at the same time to birds
J35 and J36, as controls. All injections were made into left breast muscle.
J36 became positive September 4 and died September 7 of severe malaria.
(J35 died August 30, negative.)
August 27, and September 1, 3, 5, 8, 10, 12, 15, 17, 18, and 20. Daily
blood smears from J 5 negative. (Each smear searched for 30 minutes).
September 17. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from J5 injected into J44, which had
negative blood smears September 17 to October 6. J44 became positive
263774 i
Fia. 5. Bird J5.
354 The Philippine Journal of Science 1931
October 7, which was five days after injection from 54R, which was known
to be positive.
September 17. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird J38, known to be infective, in-
jected into J5.
September 22 and 23, J 5 +; September 25 and 26, J 5 ++■++:+;
September 29, J5 -f ; October 3 and 20, J5 0.
Protocol 5. Bird J6.
August 23, 1930. Blood smear from J6 negative (30 minutes).
August 23 to 29. J6 received 0.00016 gram plasmochin simplex by
intramuscular injection about 10 a. m. each day into right breast.
August 27. J6 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U73, known to be infective.
An equal amount of the same mixture was given at the same time to birds
J35 and J36, as controls. All injections were made into left breast muscle.
J36 became positive September 4 and died September 7 of severe malaria.
(J 35 died August 30, negative.)
August 27, September 2, 4, 6, 9, 11, 13, 15, 17, 18, 20, 22, 24, 26, and 29,
and October 2 and 6. Daily blood smears from J 6 negative. (Each smear
searched for 30 minutes.)
September 17. Three-tenths cubic centimeter of physiologic saline solu-
tion mixed with 5 to 7 drops of blood from J6 injected into J45, which
had negative blood smears September 17 to October 4. J45 became posi-
tive October 6, four days after injection from 54R, which was known to be
positive.
October 2. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from J6 injected into J46, which shad
negative blood smears October 2 to 16. J46 became positive October 20,
which was five days after injection from J43, which was known to be
positive.
October 2. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird 54R, known to be infective,
injected into J 6.
October 7 to 9, J 6 +; October 10 and 11, J 6 + + ; October 13, J6
+ + + ; October 14, J 6 + + ; October 15, J 6 + + + + ; October 16, 17, and
18, J6 + + ■+ + + ; October 20, J 6 died.
Protocol 6. Bird J 7.
August 23, 1930. Blood smear from J? negative (30 minutes).
August 23 to 29. J7 received 0.00016 gram plasmochin simplex by intra-
muscular injection about 10 a. m. each day into right breast.
August 28. J7 received 0.3 cubic centimeter physiologic saline solution
containing 5 to 7 drops of blood from bird U37 known to be infective.
An equal amount of the same mixture was given at the same time to birds
J37 and J38 as controls. All injections were made into left breast
muscle. J38 became positive September 4. (J37 died August 30.)
August 28, September 3, 5, 8, 10, 12, 15, 17, 18, 20, 22, 24, 26, 29, and
October 2 and 6. Daily blood smears from J 7 negative. (Each smear
searched for 30 minutes.)
46, 3
Russell: Avian Malaria Studies, II
355
Fig. 6. Bird J6.
October 2. Three-tenths cubic centimeter of physiologic saline solution
mixed with 5 to 7 drops of blood from J 7 injected into J48, which had
negative blood smears October 2 to 22. J48 became positive October 23,
which was seven days after injection from J43, which was known to be
positive.
October 2. Three-tenths cubic centimeter physiologic saline solution
mixed with 5 to 7 drops of blood from bird 54R, known to be infective,
injected into J7.
October 7 to 10, J? + ; October 11, J7, + -f ; October 13, J7 + + + + + ;
October 14, J7 + + +; October 15, 37 + + ; October 16-22, J7 +.
Protocol 7. Bird J 8.
August 23, 1930. Blood smear from J8 negative (30 minutes).
August 23 to 29. J8 received 0.00016 gram plasmochin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
356
The Philippine Journal of Science
mi
Fig. 7. Bird J 7.
August 28. J8 received 0.3 cubic centimeter physiologic saline solu-
tion containing 5 to 7 drops of blood from bird U37, known to be infective.
An equal amount of the same mixture was given at the same time to
birds J37 and J38, as controls. AH injections were made into left breast
muscle. J38 became positive September 4. (J37 died August 30.)
August 28, September 1, 4, 6, 9. Daily blood smears from J 8 negative.
(Each smear searched for 30 minutes.)
September 9. J8 died.
Protocol 8. Bird J9.
August 23, 1930. Blood smear from 39 negative (30 minutes).
August 23 to 29. J9 received 0.00016 gram plasmochin simplex by intra-
muscular injection about 10 a. m. each day into right breast.
August 29. J 9 received at 3 p. m. 0.3 cubic centimeter physiologic sa-
line solution containing 5 to 7 drops of blood from bird U22, known to be
46,3 Russell: Avian Malaria Studies, II 357
infective. An equal amount of the same mixture was given at the same
time to birds J39 and J40, as controls. All injections were made into
left breast muscle. JS9 became positive September 3. J40 became po-
sitive September 4.
August 29, September 3, 5. Daily blood smears from J9 negative.
(Each smear searched for 30 minutes.)
September 8 to 10. J 9 +; September 12, J 9 ++'+; September 12,
J9 died.
Protocol 9. Bird J10.
August 23, 1930. Blood smear from JlO negative (30 minutes).
August 23 to 29. JlO received 0.00016 gram plasmochin simplex by in-
tramuscular injection about 10 a. m. each day into right breast.
August 29. JlO received at 3 p. m. 0.3 cubic centimeter physiologic
saline solution containing 5 to 7 drops of blood from bird U22, known to
be infective. An equal amount of the same mixture was given at the
same time to birds J39 and J40, as controls. |A11 injections were made into
left breast muscle. J39 became positive September 3. J40 became po-
sitive September 4.
August 29 and September 4 to 6. Daily blood smears from JlO nega-
tive. (Each smear of blood searched for 30 minutes.)
September 9 to 13, JlO -f ; September 15, JlO + + ; September 17 and
20, JlO +; September 25, JlO 0; October 20, JlO +.
FOURTH EXPERIMENT — QUININE SERIES
(AUGUST 25 TO OCTOBER 20, 1930)
DRUG
The drug used in these experiments was quinine dihydrochlo-
ride Lilly sold as a sterilized solution for intramuscular use.
It was purchased at a local pharmacy in boxes of 12 ampoules of
1 cubic centimeter each. According to the label each ampoule
contained 0.25 gram of quinine dihydroehloride in 1 cubic centi-
meter solution. This solution taken from the ampoules was
diluted for this experiment with distilled water, and an at-
tempt was made to determine a dose that would not produce
marked symptoms of drug toxaemia in the birds.
ManwelK2) found the minimum lethal dose for quinine to be
0.006 gram for a bird of average weight and the minimum lethal
dose of plasmochin to be 0.001 gram. His birds had an average
weight of 16.5 grams. The birds used in the experiments re-
ported in this and the first paper of the series averaged for
100 birds 15.98 grams in weight. Manwell(2) in his therapeu-
tic studies used doses of 0.000132 gram of plasmochin dissolved
in 0.1 gram of solution and 0.00075 gram of quinine (salt not
specified) in 0.00075 gram of solution. This dose of quinine he
subsequently increased to 0.001 gram in 0.1 gram of solution,
358 The Philippine Journal of Science iusi
It should be noted that in ManwelPs work the drugs were given
orally by oesophageal tube, whereas in the experiments here re-
ported the drugs — quinine dihydrochloride and plasmochin sim-
plex— were given intramuscularly.
The dose of plasmochin as already discussed in the experi-
ments here reported was 0.00016 to 0.0002 gram given in 0.1
cubic centimeter amounts.
The dose of quinine dihydrochloride finally determined as one
which would not give symptoms was 0.0005 gram given in 0.05
cubic centimeter amounts. To each ampoule of the solution as
purchased 24 cubic centimeters of distilled water were added
and 0.05 cubic centimeter of the resulting solution constituted
a dose. (Consult also Sergent.(4)) There was little or no ne-
crosis of muscle at the site of injection. Care was taken to
make deep injections well forward. The mortality in the 10-day
period beginning with the first injection of quinine was 5 per
cent in the birds receiving quinine and 40 per cent in the con-
trols which received no drugs. (See Table 1.)
PROCEDURE
Twenty canaries, Jll to J30, were each given an intramuscular
injection of quinine dihydrochloride, 0.0005 gram, each morn-
ing for seven days at about 10 a. m. into the right breast
muscle. Four birds, Jll, J12, J13, and J14, were inoculated
with infected blood from bird U90 into the left breast muscle
at 3 p. m. of the third day. In a similar way J15, J16, J17,
and J18 were inoculated from bird U91 on the fourth day;
J19, J20, J21, and J22 from bird U73 on the fifth day; J23,
J24, J25, and J26 from bird U37 on the sixth day; and J27,
J28, J29, and J30 from bird U22 on the seventh day. It will
be noted by reference to the protocols of the plasmochin series
in this paper that the same donor bird was used each day for
the two birds being protected by plasmochin and the four in
which an attempt was being made to protect by quinine.
The injections were all made with blood-saline mixture taken
from the same vial and given in the same amount — 0.3 cubic cen-
timeter— as described in the first paper. The control birds on
the third day were J31 and J32 ; on the fourth day J33 and J34 ;
on the fifth day J35 and J36 ; on the sixth day J37 and J38 ;
on the seventh day J39 and J40. These birds were also infected
from the same corresponding vials using the same blood-saline
mixture in the same amount given in the same way. The same
46,3
Russell: Avian Malaria Studies, II
359
controls, therefore, served for both the quinine and the plasmo-
ehin series and these two series in turn acted as controls to
each other.
RESULTS
It will not be necessary to give detailed protocols of each bird
receiving quinine because, with the exception of birds J19 and
J30, which died five days after inoculation, and controls J35 and
J37, which also died soon after inoculation, all of the controls
and all of the birds that had been given quinine as a preventive
became infected. Quinine failed completely to prevent infec-
tion in every case. See Table 3 for detailed results of blood
examinations in these birds and contrast them with the negative
results in the plasmochin series in all but the two birds infected
on the last day of their series of plasmochin injections.
Table 3. — Quinine series.
Number of bird.
Date inoculated,
1930.
Parasites first
seen, 1930.
Number of
days 4 or 5
plus.
Date of death.
Jll
August 25
do
do
do
August 26___
do
September 1
do
September 2
September 1
September 3
do
1
1
0
0
0
2
4
2
September 8, 1930.
October 17, 1930.
September 23, 1930.
October 6, 1930.
September 3, 1930.
Alive February 18, 1931.
September 15, 1930.
Alive February 18, 1931.
August 27, 1930.
September 13, 1930.
Alive February 18, 1931.
September 12, 1930.
September 6, 1930.
Do.
Do.
Alive February 18, 1931.
Do.
Do.
September 11, 1930.
September 3, 1930.
J12
J13
J14
J15
J16
J17
J18
J19
do
do
August 27 __. .
September 2
do
J20
J21
J22
do
do
_ do
September 3
September 4
do
3
0
3
0
0
0
1
0
0
0
J23
August 28
do
September 5
do
J24
J25
J26
J27
do
do
August 29
do ._._
September 6
September 4
September 5
do
J28
J29
J30
do
do..
September 4
The fact that the average prepatent period in the birds given
quinine was 7.4 days and in the controls was 6.5 days is of
doubtful significance because no attempt was made to adjust
the quantity of the infective blood-saline inoculum to the size
of the birds. The same dose of 0.3 cubic centimeter was given
in every case as noted in the first paper of this series. (1)
360 The Philippine Journal of Science
Boyd (3) found that the prepatent period is a function of the
size of the inoculum.
SUMMARY
Two experiments are reported in which attempts were made
to prevent the infection of canaries following needle inocula-
tions with P. cathemerium (Hartman, 1927). In the first ex-
periment plasmochin simplex and in the second experiment
quinine dihydrochloride was given in daily intramuscular doses
for a week. Infection was attempted in some birds on the
third day; in others on the fourth, or fifth, or sixth, or seventh
day. The birds that had had plasmochin for seven days and
were infected on the seventh day, five hours after their last dose
of plasmochin, became infected, as did all of the birds receiving
quinine. When infection was attempted on the third, fourth,
fifth, and sixth days of the plasmochin series the attempt in-
variably failed.
CONCLUSION
1. It is concluded that the infection of a canary by experi-
mental needle inoculation with P. cathemerium (Hartman,
1927) can be prevented by intramuscular injections of plasmo-
chin simplex in daily doses of 0.00016 gram, provided that the
bird receives at least one dose of plasmochin subsequent to
receiving the infective inoculum.
2. It is concluded that the protective power of plasmochin in
these needle inoculations is transitory and does not persist as
long as five hours. (It may, therefore, be reasonable to con-
clude as a corollary that the protective power of plasmochin in
experimentally inoculated malaria in birds is more therapeutic
than preventive.)
3. Finally, it is concluded that quinine dihydrochloride in daily
intramuscular doses of 0.0005 gram will not protect birds from
experimental needle inoculations with P. cathemerium (Hart-
man, 1927).
BIBLIOGRAPHY
1. Russell, P. F. Avian malaria studies, I. Philip. Journ. Sci. (Pre-
cedes this paper.)
2. Manwell, R. D. Am. Journ. Trop. Med. 10 (1930) 379-406.
3. Boyd, G. H. Am. Journ. Hyg. 6 (1926) 173-195.
4. Sergent, Et. and Ed. Ann. d. Tlnstitut Pasteur 35 (1921) 1-17.
Fig.
ILLUSTRATIONS
TEXT
FIGURES
1.
Plasmochin
simplex,
a prophylactic
Fourth experiment.
2.
Bird Jl.
3.
Bird J2.
4.
Bird J3.
5.
Bird J5.
6.
Bird J6.
7.
Bird J7.
361
MALARIA TRANSMISSION IN THE PHILIPPINES, V
ON THE MATURATION OF THE OVA OF ANOPHELES FUNESTUS GILES 1
By C. Manalang
Of the Philippine Health Service, Manila
ONE TEXT FIGURE
In the 7th Congress, Far Eastern Association of Tropical
Medicine, held at Calcutta (December, 1927), James, Nicol, and
Shute2 reported higher mortality of Anopheles maculipennis
Mg. in some months than in others, as observed during a period
of three and one-half years on forty-one batches of artificially
infected mosquitoes in England. In June less than 2 per cent
(should be less than 20 per cent or 17 per cent, an error in
the decimal; see their table) survived to be infective, while
in October at least 50 per cent would be available. By field
observation they explained this to be due to the process of egg
maturation and deposition, which caused high mortality. Their
observations agreed with those of Swellengrebel in Holland 3
in regard to the period of egg maturation and oviposition of
maculipennis (spring and summer) when no positive mosquitoes
were found, and the autumn and winter months when the ova
were not developed but with natural malaria infection in the
mosquitoes. Their conclusions are:
The lessons of these observations from the point of view of the spread
of malaria seem to be (1) that in the future we must endeavor to corre-
late the seasonal incidence of primary malaria, not with the seasonal prev-
alence of mosquitoes concerned but with the seasonal prevalence of
the individuals which live long enough to be transmitters. In June, there
may be an enormous number of adult maculipennis in a malarious place
but if we know that during that month less than 2% [should be less than
20 per cent] live long enough to become transmitters of the disease, their
abundance is not so important. Obviously, it is much less important than
a smaller abundance in August or September; the simple calculation from
1From the field laboratory, division of malaria control, Philippine Health
Service, Tungkong Manga, Bulacan.
2 Trans. 7th Congress F. E. A. T. M. 2 (1927) 712-717.
8 Malaria in the Kingdom of the Netherlands. Report to the malaria sub-
committee of the Health Committee of the League of Nations (1927).
363
364 The Philippine Journal of Science i9si
our figures that 100 mosquitoes in September are equal in importance to
3,000 in June [this should be 300, see their table] does not by any means
express the true difference because the September mosquitoes will live
several months while the June mosquitoes will live at the most only a few
weeks; (2) If the process of egg maturation and oviposition is such an
important cause of death that it almost entirely prevents the transmission
of malaria by anopheles during the months of its occurrence, the number
of broods that each species has in different localities and periods of the
year during which maturation of eggs and oviposition occurs ought to be
worked out much more carefully than has hitherto been attempted in many
places. The results may provide a clue to the explanations of some ob-
servations on malarial incidence which are at present obscure.
Boyd,4 using a standard measure of A. qttadrimaculatus Say
density (mosquitoes caught per man-hour search) noticed ex-
plosive increases in density, particularly those of the males, and
concluded that in the latitude observed in southwestern Georgia
this species may have from eight to ten annual generations, or
a monthly generation, excepting in January and February.
His records of dissection led him to conclude that "(a) The
occasions when gland infections have been found have been pre-
ceded by the detection of stomach infections, and (b) Referring
to the hypothetical brood curves, the stomach infections occurred
at a period when the density of a generation has been at its
maximum, while gland infections were found at the period when
the brood was frankly in decline." As to ovarian development
he found nulliparous females most abundant when the brood is
on its upward phase and multiparous females encountered when
the brood is on the wane. Boyd and Weathersbee,5 observing
along the coast of North Carolina (36° north latitude) found that
in qwadrimaculatus and punctipennis "gravid females were rel-
atively high in number in the early winter and until oviposi-
tion began in January and February. Digestion of blood and
development of the ovaries proceeds slowly thruout winter, the
gravid females tending to withhold oviposition until the tem-
perature is favorable."
Boyd6 in a more recent article (p. 457) says:
No gravid females of quadrimaculatus were found during winter period.
This shows that during the winter the distended abdomens of stage e fe-
males (Grassi and Sella) are to be attributed to an hypertrophied fat body.
No instance of reproductive activity during the coldest winter month was
4 Am. Journ. Hyg. 7 (1927) 264-275.
5 Am. Journ. Hyg. 9 (1929) 682-694.
6 Am. Journ. Hyg. 12 (1930) 449-466.
46,3
Manalang: Malaria Transmission, V
365
found. The statement in Boyd and Weathersbee (8) that "the majority of
A. quadrimaculatus found in unoccupied places were gravid" is incorrect
in the light of this information. It appears, therefore, that the1 imagines
of quadrimaculatus encountered during the fall or winter periods are
destined for, or are actually undergoing hibernation.
His table (p. 463) of seasonal distribution of infected speci-
mens for 1926, 1927, and 1928 shows that the infections were
found between June and October.
King's 7 twelve months' observation on natural malaria in-
fection in the vicinity of Mound, Louisiana, records no infections
in 1,375 quadrimaculatus caught in winter (November to April),
but 12, or 1.96 per cent, of the 611 caught in summer (May to
October) were infected.
The data for the present paper were collected from Septem-
ber, 1927, to August, 1929, in the La Mesa and South Portal
camps of the Novaliches water project.
Table 1. — Anopheles funestus caught in two years at La Mesa and South
Portal giving percentages of insects with matured ova and the rates
of infection.*
Month.
January. _.
February -
March
April
May
June
July
August
September
October __.
November.
December.
Mosquitoes caught and
dissected.
Total.
932
327
878
668
686
782
883
571
1,058
1,190
767
803
With mature
ova.
Stomach positive.
51
25
55
115
71
103
98
116
48
105
33
47
Percent.
5.4
7,
6
17,
10.
13
11
20
.6
.2
.2
.3
.1
.1
.3
4.5
8.8
4.4
5.8
Per cent
1.2
1.8
1.7
0.6
2.6
1.0
2.6
3.0
0.7
1.0
1.0
1.7
Salivary gland
positive.
Percent.
0.8
0.9
1.6
1.3
2.8
1.4
1.1
1.2
0.7
0.1
0.5
1.1
a Matured ova in a positive mosquito were found in only one case (salivary gland)
among almost 300 positives observed from six areas. Engorged stomach with oocyst was
not uncommon. In 15 cases both stomach and salivary glands were infected.
The table gives the monthly number of A. funestus Giles dis-
sected, the number and percentage with ova (float and chitinous
structures formed), and the stomach and gland infections. The
7 Am. Journ. Hyg. 1 (1921) 35-39.
366
The Philippine Journal of Science
1931
graph (fig. 1) shows a rise in mosquito infection coinciding with
the months with the high rate of matured ova.
COMMENTS
The findings on funestus are, therefore, diametrically opposed
to those of James in England and Swellengrebel in Holland that
oviposition of maculipennis prevented malaria transmission due
to shortening of its life.
The findings of quadrimaculatus by American investigators
also do not agree with those of the Europeans.
20
19-
18
17
16
15-
14
13-
IZ^
IH
10-
9
8
+* 1
C 6
8 S
\
\
/ \
A
/ \
/ \
y
\
/ \
/
\
/ x
/
\
tJ
1
\
,/N
. /
c</
\ y
sy
/
V
\
/
\
/
\
A
/
\
/ \
\/
\ /
' \
\/
\
v
\^
°- 3-
2-
i-
s
alive
;py^
land
\
\
-Sto
mac
h
/
/
/
'*"*-«—-
1
\
\
\
- -r-r ^
/
i
0
V
X
*s. ^
.>*•
(O
Q_
<
(O
CM
C
■ 3
<
o
O
o
21
CD
Fig. 1. Percentages of AiiopTieZes funestus Giles with ova and with stomach and salivary-
gland infections. La Mesa and South Portal camps, Novaliches water project, Luzon,
September, 1927, to August, 1929.
The coincidence of higher malaria-infection rates in funestus
with the higher rate of the females with matured ova indicates
that this species not only survives oviposition but that the
period with the maximum rate of matured ova is the period of
maximum malaria transmission. This seems logical on biol-
ogical grounds, and as indicated in my preceding article, the
rates of infection in funestus rose with rainfall, temperature,
and humidity. These meteorological factors undoubtedly ac-
46,3 Manalang: Malaria Transmission, V 367
celerate egg development and induce the mosquito to seek blood
for the maturation of the eggs; consequently to bite man and
become infected with gametes. When the ova mature and are
deposited, more immature ova develop with a desire for human
blood, so that with sporozoites in the salivary glands the mos-
quito bites man and infects him.
SUMMARY
1. Data on malaria infection and maturation of the ova of
Anopheles funestus in La Mesa and South Portal camps of the
Novaliches water project, Luzon, show higher rates of the
former during the period of high rates of the latter, a finding
opposite to those of James in England and Swellengrebel in
Holland, who held that egg maturation and oviposition prevent
malaria transmission due to the shortened life of A. maculi-
pennis.
2. Findings on A, quadrimaculatus by American workers
agree with the findings in the Philippines.
3. The findings on A. funestus seem logical on biological
grounds.
ILLUSTRATION
Text fig. 1. Percentages of Anopheles funestus Giles with ova and with
stomach and salivary-gland infections. La Mesa and South
Portal camps, Novaliches water project, Luzon, September,
1927, to August, 1929.
263774 5 369
MALARIA TRANSMISSION IN THE PHILIPPINES, VI
THE DARK-NIGHT FACTOR *
By C. Manalang
Of the Philippine Health Service, Manila
It is a general practice among anopheles-mosquito experi-
menters to cover the mosquito cage with black cloth if it is
desired that the mosquitoes bite during the day or to expose the
patient to them at night.
During December, 1927, and May and June, 1928, it was no-
ticed on Anopheles funestus Giles dissection records from South
Portal camp of the Novaliches water project, that malaria in-
fection coincided with the new-moon period of the month, or a
few days before or after the new moon.
To determine the relation between the positive catches and
the moon periods, the calendar was divided into two periods of
fifteen days each; namely, the full-moon period (seven days
before and seven days after the full moon) and the new-moon
period (seven days before and seven days after the new moon).
In January, 1928, these two periods had an interval of one day,
the fifteenth, while the new-moon period in the last half of
the month overlapped one day with the full-moon period of
February, on January 30; in March, the interval was on the
fourteenth and the overlap on the twenty-ninth; in April, the
interval was three days (the thirteenth, fourteenth, and fif-
teenth) and the overlap was two days, the twenty-ninth and
thirtieth ; in May, the overlap was on the twelfth ; in June, the
interval was on the twenty-sixth; in July, the overlap was on
the tenth; in August, the overlap was on the eighth, with the
interval on the twenty-third ; in September, the overlap was on
the seventh; in October the overlap was on the sixth; and in
November, the overlap was on the fifth and the interval on the
twenty-first.
1 From the field laboratory, division of malaria control, Philippine Health
Service, Tungkong Manga, Bulacan.
371
372 The Philippine Journal of Science 1931
The mosquito data in South Portal from December, 1927, to
December, 1928, inclusive, are used in the present paper. These
data contain the largest number of funestus infections registered
in all the camps studied. Dissection findings were entered on
the date of capture and not on the date of dissection. They
were kept for several days after capture to allow positive
blood meals to develop.2 Positives falling on the overlapping
dates were credited to the period before it. Two positives were
thus credited to the new-moon period and three to the full-
moon period. No positives were found in the intervening dates
between the two periods except one on April 13, the eighth day
after the full moon on April 5, which was credited to the full-
moon period, there being an interval of three days.
Table 1 shows the distribution of positives in these periods
and shows the influence of the darkness of the new-moon period
on the number of infected funestus.
The darkness of the night was not recorded at the La Mesa
camp, so it is not possible to correlate the mosquito findings
during the dry months, December to April. The rain observa-
tion chart, however, indicated rain-gauge readings at 6 a. m.
and 2 p. m. and noted the beginning and duration of the rai^n
during the day or night. There was no record of the rainless
but dark (cloudy) nights during the full-moon periods. In-
vestigation of the rainy nights during the rainy months showed
that thirty-two positives of the total seventy-two caught during
the full-moon period, as shown in Table 1, were caught during
twenty-one rainy nights from May 4 to November 27. Since
these nights were usually dark, the relation of funestus infec-
tion and dark night is very clear.
Table 2 shows the number of infected funestus caught during
dark and bright nights. Dark cloudy nights from December to
April (dry season) and dark rainless nights during the full-
moon period are not known.
By including the period, December, 1927, to April, 1928, when
the darkness or brightness of the night was based only on the
2 In an unsuccessful attempt to compare laboratory infectivity of dif-
ferent anopheline species, ten carriers were bitten by 1,157 mosquitoes with
the following result: One vagus with two matured oocysts died on the eighth
day after the infective meal; one funestus with three matured cysts on
the fourth day; one karwari with four matured oocysts on the sixth day;
and one ludlowi with twenty-two matured oocysts on the seventh or ninth
day after the infective meal.
46,3
Manalang: Malaria Transmission, VI
373
new- and full-moon periods on the calendar, 80 per cent of the
infected funestus were caught in the dark-night period. Ex-
cluding the December to April data (dry months), 87 per cent
of the positives were caught during the known dark nights
(with or without rain) and 13 per cent during the rainless
nights, some of which, however, might have been bright or
cloudy. Therefore, a susceptible has at least four times more
chance of contracting malaria during the dark than during
the bright nights, or 80 per cent of the infections are acquired
during the dark nights. The moon has apparently no influence.
Table 1. — Malaria in funestus distributed in the two moon periods.
Month.
Number of positives.
New-moon
period.
Full-moon
period.
December, 1927_
6
7
3
7
7
21
12
23
14
10
8
1
7
0
7
4
6
4
10
6
4
9
11
3
2
7
January, 1928__
February
March
April
May
June _ __
July__ _
August _
September
October _ __
November
December
Total*
126
63.6
72
36.4
Per cent __
* Positive stomachs, 113 ; positive salivary glands,
COMMENTS
85.
Table 3 shows the number of mosquitoes caught, the number
of days employed in catching, and the average daily catch for
the dark and bright nights (by calendar) and shows an average
of twenty-three for the former and twenty for the latter. The
twenty-one rainy nights of the full-moon periods from May to
November gave a daily average of twenty-three funestus. So
there was no increase in density during the dark nights, whether
rainy or not. Larger numbers of them have been observed to
enter the trap between 10 p. m. and 2 a. m. than at other times.
374
The Philippine Journal of Science
1931
Table 2. — Number of infected funestus caught during dark and bright
nights .*
Month
Number of positives.
Dark-
night
period.
Bright-
night
period.
December, 1927_.
January, 1928__.
February
March
April
May
June
July
August
September
October
November
December
Total
Per cent.
6
7
3
7
7
24
16
23
23
19
11
158
80
40
20
* The number of cloudy or dark nights during full-moon periods of dry months, from
December, 1927, to April, 1928, inclusive, is not known. The number of rainless but dark
(cloudy) nights during the full-moon period is also not known.
Anopheles funestus, like the other anophelines, undoubtedly
prefers to bite at night, as observed in experimental work. The
camp people probably retire earlier during dark or rainy nights
and give the mosquitoes an undisturbed chance to bite.
Culicines and some anophelines (vagus and ludlowi) are at-
tracted by yellowish lantern light. At one of the Tungkong
Manga laboratory traps, A. philippinensis came by the hundreds
on two nights, attracted by the bright white light from an Alad-
din lamp. Funestus were caught in South Portal traps with or
without lantern light, so that it could not be said that a light
in the trap guided them during dark nights. In the laboratory
traps at Tungkong Manga, about 400 funestus were caught dur-
ing the past ten months without using light.
The influence of bright or colored lights has not been tested.
The study of illumination and transmission may give some clue
to the factors causing prevalence of malaria in the rural,
newly developed, out-of-the-way districts in the Philippines.
46,3
Manalang: Malaria Transmission, VI
375
Table 3. — Comparative density of mosquitoes between the two moon
periods.
Month.
December, 1927_.
January, 1928__.
February __.
March__
April
May
June
July „
August
September
October
November
December
Total
Daily average.
New moon.
Number
caught.
157
396
83
306
332
286
239
357
197
270
340
99
134
3,196
Number
of days.
140
Daily
average.
31
44
10
33
26
22
20
30
18
22
26
12
12
23
Full moon.
Number
caught.
17
355
140
205
171
210
235
207
371
109
219
208
169
52
2,668
Number
of days.
1
11
7
10
8
10
10
10
12
12
13
12
10
8
134
Daily
average.
17
33
20
20
21
21
23
21
31
9
17
17
17
7
SUMMARY
1. Data collected during thirteen months of study of natural
malaria infection of A. funestus in South Portal camp of the
Novaliches water project show that at least 87 per cent of
those infected were caught during the dark nights. Malaria
transmission in the locality observed was at least four times
more during the dark than during the bright nights, and the
chance of contracting the disease was eighty times out of a
hundred during the dark nights.
CAUSES OF IRRITATION UPON INJECTION OF IODIZED
ETHYL ESTERS OF HYDNOCARPUS-GROUP OILS 1
By Howard Irving Cole
Chief Chemist, Culion Leper Colony, Philippine Health Service
At the recent conference of leprologists held under the aus-
pices of the Leonard Wood Memorial for the Eradication of
Leprosy, the majority of the delegates agreed that the ethyl
esters of hydnocarpus-group oils are among the most active
drugs at present available in the treatment of leprosy. Several
leprologists stated that, by their methods of preparation, a non-
irritating oil was essential for the production of esters of
low-irritant quality. It is not always possible, however, for
institutions situated many thousands of miles from the sources
of supply to obtain oil that is fresh and nonirritating. It, there-
fore, becomes of prime importance that a method be devised by
which even intensely irritating oils may be utilized for making
relatively nonirritating ethyl esters. In order to accomplish
this, it is necessary to take into consideration all the factors that
may cause irritation and vary the method of preparation of the
esters accordingly.
It has previously been found 2 that irritant properties of ethyl
esters may be reduced by (1) elimination of free fatty acids;
(2) elimination of decomposition products due to heating or
chemical treatment; (3) elimination of volatile and nonvolatile
impurities ; and (4) addition of 0.5 per cent iodine.
Free fatty acids can be reduced to a minimum (less than
0.2 per cent) by careful neutralization with sodium hydroxide
and very thorough washing. Free fatty acids are undoubtedly
one of the main causes of irritation. To be certain that the
amount present is less than 0.2 per cent, titration with tenth
normal alkali should be made a part of the routine procedure for
the preparation of the ethyl esters.
1 Published with the approval of the Director of Health.
» Cole, Philip. Journ. Sci. 40 (1929) 503.
377
378 The Philippine Journal of Science mi
Decomposition products, formed upon heating or strong chem-
ical treatment, and other volatile impurities can be largely
blown out by steam.
If the esters are distilled, nonvolatile impurities and decompo-
sition products remain in the still as residue. The distilled
esters are decidedly more limpid than the undistilled, due prob-
ably to the fact that the latter contain some unchanged oil.
Addition of 0.5 per cent iodine markedly reduces the irritant
effect of the ethyl esters providing that the method of iodization
described below is strictly followed, in which case, the iodine
is in the combined form. The presence of free iodine causes
irritation.
The inherent irritant quality due to the configuration of the
molecule of the compound is, of course, not removable without
changing the compound itself. It has been shown that a syn-
thetic compound similar to ethyl hydnocarpate, pure dinormal
heptyl ethyl acetate,3 is even more irritating than ethyl hydno-
carpate, while the glyceryl ester of this synthetic compound
(corresponding to a natural oil) is bland. This would indicate
that part of the irritant effect of the ethyl esters is associated
with the ethyl radicle.
A method of preparing hydnocarpus ethyl esters of a standard
low-irritant quality, no matter how irritating the original oil
may be, has already been described.4 Since the publication of
this process, however, continued experimentation has thrown
further light upon the causes of irritation and their prevention.
RELATION BETWEEN IRRITATION AND TYPE AND SHAPE OF
CONTAINER
Our standard method 5 for iodizing ethyl esters is as follows :
Fifteen liters of the purified esters are heated in a 20-liter enameled
or stainless steel kettle to 140° C. The esters must be thoroughly dried
before iodine is added since, if water is present, it effects by catalysis
the hydrolysis of several per cent of the esters. If the filtered esters are
clear, the heating to 140° C. before adding the iodine will drive off all
dissolved water. Seventy-five grams of chemically pure resublimed iodine
are added with stirring. The temperature immediately rises to 150° C,
at which point it is maintained for exactly thirty minutes, the liquid
being stirred occasionally. After cooling, the iodized esters are filtered
into bottles (250 cubic centimeters capacity) and sterilized for one hour
* Private communication from C. B. Lara; drug prepared by Roger Adams.
4 Cole, Philip. Journ. Sci. 40 (1929) 503.
•Loc. cit
46'3 Cole: Catises of Irritation 379
in an oven at 150° C. The temperature of the contents of the bottles
reaches in this time 110° C.
Since this method of iodization was adopted, more than 3,000
liters of ethyl esters (200 lots of 15 liters each) have been
iodized and used with practically no complaints of excessive
irritation. Smaller institutions, however, might desire to make
smaller lots of esters. It was found, in certain recent experi-
mental work, that when the drug was iodized in 2-liter lots
instead of the standard 15-liter batches, it was more irritating
than usual, although the standard method was carefully followed,
except for the fact that these lots were heated in tall, 3-liter
glass beakers instead of low, stainless steel or enameled kettles.
Experiments were then made using enameled beakers instead of
glass beakers, but the drug so prepared was no less irritating.
The form of the container was changed from a tall (beaker)
type to a shallow (pan) type.6 The preparations were equally
irritating provided that the time of heating to 150° C. and time
of cooling to room temperature of these two types were the same.
In ordinary practice, however, with the low form (pan type),
the drug heats more quickly and cools more rapidly than with
the tall form. The longer time necessary to heat and cool the
contents of the tall type of container corresponds to overheating
of the drug, and we already know that overheating results in
an irritant product. This rapid heating and cooling is evidently
preferable, for the product in this case was less irritant and, in
fact, entirely comparable with that produced by the standard
method for 15-liter lots.
EFFECT OF STIRRING ON IODIZATION
In order to determine whether stirring during the iodization
is beneficial or otherwise, two batches of esters were iodized at
the same time under identical conditions, except that one lot
was gently stirred only during the addition of the iodine, while
the otfter one was vigorously stirred by means of a motor stirrer
during the entire heating. The time of heating to 140° C. (15
minutes), the time of heating with iodine at 150° C. (30 min-
utes), and the time necessary to cool to 40° C. (2 hours) were
kept constant for both lots by regulating the heat input. No
• It is assumed that the container will be more than half filled. The depth
of the liquid would then be greater than the diameter in the tall type
and should be not much more than half the diameter in the shallow type.
Missing
Page
Missing
Page
382
The Philippine Journal of Science
Table 2. — Effect of light, heat, and air on color and irritation of
iodized ethyl esters.
Before exposure
Exposed to direct sunlight, 10 hours
Exposed to direct sunlight, 14 hours _
Exposed to direct sunlight, 14 hours plus 16 days in dark_.
Exposed in dark, 10 days at 30° C _
Exposed in dark, 14 hours at 50° C
Exposed in dark, 10 days at 50° C
Color in millimeters
equivalent to 20 mil-
limeters of standard.
Filled
stoppered
bottles.
22
22
22
22
22
22
22
Open
beakers.
22
♦35
*41
*60
24
28
*60
* Products marked thus were found to be very irritating upon injection.
Occasional stirring during iodization of the esters is probably
beneficial. Continuous vigorous stirring is not necessary.
Experiments show that color comparison cannot be utilized as
control in the production of standard relatively nonirritating
iodized ethyl esters; time and temperature of heating of the
esters with the iodine must be used as the basis of control.
Sunlight or heat in the presence of air soon changes the iodized
esters in such a way as to yield an extremely irritating product.
This deterioration is accompanied by increased clarity and
change in color to reddish brown.
The author wishes to gratefully acknowledge his indebtedness
to Dr. C. B. Lara and the medical staff at Culion for performing
the irritation tests mentioned in this article.
DIE BRENTHIDEN DER PHILIPPINEN-INSELN
Von R. Kleine
Stettin, Deutschland
SECHSZEHN KARTEN
Der Catalogus Coleopterorum Junk-Schenklings, ed. 1, enthalt
nur 8 Arten von den Philippinen, davon sind 2 synonym so dass
tatsachlich nur 6 Arten bekannt waren. Heute betragt der Bes-
tand 124 Arten.
In letzten Jahrzehnt sind umfangreiche Ausbeuten von den
Inseln gekommen. Namentlich hat der leider viel zu fruh ver-
storbeaie Charles Fuller Baker ein gewaltiges Material an Indi-
viduen zusammengebracht, das zum kleineren Teil von Prof.
Heller, Dresden, spater auf Hellers Empfehlung mir zur Bear-
beitung iiberlassen worden ist. Im Zeitraum von mehreren
Jahren habe ich zahlreiche Sendunge Bakers bearbeitet und einen
guten Einblick in die Brenthidenfauna der Philippinen tun kon-
nen. Dazu kommt noch ein Teil des Materials, das Boetteher,
der, wie bekannt, fur Moser gesammelt hat, mitgebracht hatte.
Ferner sei noch auf das in der Sammlung des Bureau of Science
zu Manila hingewiesen. Soviel darf ich wohl heute ohne Ueber-
treibung sagen: der wesentlichste Bestand der philippinischen
Brenthiden-Arten ist heute bekannt. Was noch unbearbeitet in
Museen liegt — es kann sich nur um das, friiher in Mosers Besitz
befindliche Material handeln — kann nicht mehr aufregen. Was
wir von der Brenthidenfauna der philippinischen Inselwelt wis-
sen wollen, wissen wir.
Die Arbeit soil einen rein zoogeographischen Charakter tra-
gen.
Gelegentlich der Bearbeitung der philippinischen Lyciden x
habe ich in der Einleitung zur Zoogeographie f olgendes gesagt :
Nach meinen Erfahrungen, die ich in jahrelanger Bearbeitung bei den
Brenthiden gemacht habe, sind die Philippinen unbedingt zum austro-ma-
layischen Gebiet zu rechnen. Der papuanische Einschlag, der sich nament-
lich in der Ausfarbung zeigt ist so bedeutend, dass man die Philippinen
Philip. Journ. Sci. 31 (1926) 34.
383
384 The Philippine Journal of Science 1931
als einen abgesprengten Teil Neu-Guineas ansehen konnte. Die Bezie-
hungen zu den Molukken waren sehr gering, dagegen erwies sich als sicher,
dass eine Zuwanderung aus dem orientalischen Gebiet stattgefunden hatte.
Bei den Einwanderern handelte es sich um grosse, weitverbreitete Gattung-
en, die zum Teil auf den Philippinen mit ihrer Wanderung zu Ende
gekommen waren und keinen Anschluss auf der sudostlichen Zugstrasse
iiber Celebes gefunden hatten. Zum Teil sind es Arten, die im indo-ma-
layischen Untergebiet weitverbreitet sind, also eine grosse Migrationsfahig-
keit und -geschwindigkeit besitzen. Viele der Zuwanderer wurden auf
Palawan festgestellt. Dass die Zuwanderung unbedingt iiber diese Inseln
stattgefunden haben muss ist damit nicht gesagt, ich glaube vielmehr,
dass der Zustrom im Zuge der Sulu-Inseln wenigstens ebensogross, wenn
nicht noch grosser, gewesen ist. Leider sind diese zoogeographisch so
wichtigen Inseln noch nicht exploriert.
Diese Ansicht halte ich audi heute. noch auf recht und sehranke
sie nur insoweit ein, als der Zustrom aus dem orientalischen
Gebiet beiden Brenthiden grosser ist, als aus dem austro-malay-
ishchen, eben, weil die Abtrennung von dem sich um Neu Guinea
gruppierenden Landmassiv eine vollkommenere ist, als von den
Sunda-Inseln. Wie in meiner Arbeit iiber die Lyciden, habe
ich Palawan auch hier nicht zu den Philippinen gerechnet. Sie
gehoren organisch und faunistisch zu Borneo.
DIE VERBREITUNG AUF DEN EINZELNEN INSELN
Von Marinduque, Catanduanes, Burias, Tablas, Cebu, Ticao,
Biliran und Dinagat lag mir kein Material vor. Die Werbrei-
tung auf den einzelnen Inseln war f olgende : Von den 124 Arten
waren 6 ohne nahere Fundortangabe, 188 mit sicheren Fundorten
belegten Arten f anden sich auf :
Luzon
58
Siargao
7
Polillo
4
Panay
3
Mindoro
2
Negros
26
Masbate
1
Mindanao
66
Samar
20
Basilan
15
Sibuyan
7
San Miguel
5
Leyte
9
Panaon
2
Bohol
1
Die Gattungen sind, soweit sie mehrere Arten umfassen, auf
den ganzen Archipel verbreitet. Die einzelnen Arten sind,
wenigstens nach unseren bisherigen Kenntnissen, zum teil lokal.
Bei haufigeren Arten kann man aber leicht feststellen, dass die
einzelnen Inseln wahrscheinlich keine Endemismen beherrber-
geoi, dass die Arten vielmehr auf alien Inseln zu finden sein
werden. In nachstehender Tabelle ist die Verbreitung der ein-
zelnen Arten angegeben.
Endemisch
47.6
Java
Ceylon
8.1
Andamanen
Indien
16.2
Formosa
Bengalen
20.0
Japan
Indo-China
9.0
Celebes
Malay Halbinsel
31.4
Molukken
Sumatra
34.7
Neu-Guinea
Borneo
36.3
Australien
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 385
Prozent Prozent
28.2
9.0
10.5
1.6
9.0
15.3
7.4
5.7
Danach zahlen zum Indischen Untergebiet 53.3 prozent, zum
Gebiet das sich um die grossen Sunda-Inseln und Malakka grup-
piert 139.6 prozent, zum Palaearktikum 11.6 prozent und zum
austro-malayischen und australischen Gebiet 37.4 prozent.
Vergleicht man die Zahlen in Bezug auf die mutmassliche Zu-
wanderung, so ist das Verhaltnis wie 204.5 : 37.4 oder wie
5.5 : 1.
Am iibersichtlichsten sind die Zahlen, wenn sie relativ ange-
wandt werden, wie das vorstehend geschehen ist. Da fallt zu-
nachst die hohe Zahl der Endemismen auf. Nicht weniger als
47.6 prozent kommen nur auf den Philippinen vor. Das ist
fast die Halfte aller Arten. Es muss angenommen werden, dass
sich diese Zahl nicht wesentlich verandert, denn die Brenthiden-
fauna der orientalischen Region ist so weit bekannt, dass sich
nicht viel mehr, wenigstens was die Zusammensetzung und Ver-
teilung anlangt, verandern wird.
Wie steht es nun mit dem Vergleich zu anderen Faunengebie-
ten? Ich will von einem Vergleich mit der aethiopischen, ma-
degassischen und neotropischen Region absehen, da dariiber bei
Betrachtung der Tribus gesprochen wird. Es sollen hier nur
Gebiete in Frage kommen, in denen philippinische Arten ange-
troffen worden sind. Ich fasses diese Gebiete in drei Gruppen
zusammen: 1. Die eigentliche indische, westliche. Hierzu
sind zu zahlen : Ceylon, Indien, die um die Bucht von Bengalen
liegenden Gebietsteile und Indo-China. Summiert man die rela-
tiven Zahlen so ergeben sich 53.3. Der indischen Gruppe mochte
ich : 2. Ein zentrales Gebiet entgegenstellen, das die malayische
Halbinsel, die Sunda-Inseln und die kleinen Inseln, die daran lie-
gen, einbegreift. Die Addition der relativen Zahlenwerte gibt
hier die Summe von 139.6. Als drittes kleines Gebiet soil Formo-
sa und Japan gelten, die Gesamtsumme betragt 11.6. Die ost-
lich und siidostlich des zentralen Gebietes liegenden Molukken
einschliesslich Celebes 24.3 und endlich Neu-Guinea mit Aus-
tralien und den polynesischen Inseln 13.1.
263774 6
386
The Philippine Journal of Science
1931
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46,3 Kleine: Die Brenthiden der Philippinen-Inseln 391
Ich habe in meinen zoogeographischen Studien mehrfach dar-
auf hingewiesen, dass ich zwei Ausgangszentren annehme: das
zentrale Afrika, aus welchem eine Wanderung nach Osten und
Westen stattgefunden hat und ein grosseres, zusammenhan-
gendes Landmassiv, dessen Reste in Neu-Guinea, Australien und
dem gewaltigen Inselreich, das sich von Celebes bis Tahiti hin-
zieht hinzuzurechnen ist. Von hier aus hat eine zirkumpolare
Ausbreitung stattgefunden. Auch nach Nordwesten sind die
hier hergehorigen Formen gewandert, ohne indessen den aus
Osten kommenden Zug der Familiengenossen Einhalt zu tun.
Der aus Westen kommende Wanderzug hat sich als der starkere
erwiesen. Von den 124 auf den Philippinen gefundenen Arten
gehoren 107 dem westlichen (afrikanisehen) Formenkreis an
und 17 dem ostlichen (austro-malayischen, bezw., australischen).
Die Brenthiden-Fauna der Philippinen muss also als orientalisch
angesehen werden.
Es ware noch die Frage zu erortern, ob sich auf den einzelnen
Inseln ein besonderer Typ in der Ausfarbung ausgebildet hat.
Es gibt Gebiete, zum Teil noch kleiner als die Philippinen, die
einen bestimmten Farbentyp erkennen lassen und wo es leicht
ist, die Zugehorigkeit der Art zum Gebiet festzustellen. Das
kann man hier nicht sagen. Was die allgemeine Ausfarbung an-
langt, so ist eine Tatsache allerdings sehr beachtenswert, nam-
lich, dass sich auf den Philippinen, und zwar nur dort,
Farbenkomponenten zusammengefunden haben die sonst nur in
Neu-Guinea und den angrenzenden Inseln zu finden sind:
schwarze bis blauschwarze Grundfarbe und ziegelroter Pro-
thorax. Es muss aber gleich darauf hingewiesen werden, dass
nicht nur Arten siidostlicher Provenienz davon betroffen sind,
sondern auch solche orientalischer Herkunft. Es kann also wohl
mit Recht angenommen werden, dass die Inseln noch zum Land-
massiv Neu-Guineas gehorte als die Hauptwanderung von Ost
nach West und umgekehrt bereits beendet war.
Weiter sind die Deckenzeichnunger auf den Elytren insofern
bemerkenswert, als sie die im vorigen Abschnitt geausserte An-
sich iiber die Zugehorigkeit zum alten Landmassiv Neu-Guineas
dadurch unterstatzen, dass sich eine ganz ausgesprochen Langs-
streifung bemerkbar macht. Die Orientalen haben eine entge-
gengesetzte Tendenz. Zu beachten ist die Tatsache, dass Pala-
wan nicht zum Farbungsgebiet, wie iiberhapt nicht zu den
Philippinen gehoren, sondern zu Borneo. Eine genaue Durch-
f orschung der Palawan-Inseln ware von hochstem Interesse, um
festzustellen, welche Bedeutung ihnen als Briicke zukommt.
392
The Philippine Journal of Science
1931
SYSTEMATISCHER KATALOG DER PHILIPPINISCHEN BRENTHIDEN
CALODROMINI
Genus CALODROMUS Guerin
Calodrmnus Guerin, Mag. Zool. (1832) t. 34.
CALODROMUS CRINITUS Kleine.
Calodromus crinitm Kleine, Arch. Nat. A. 10 87 (1921) 24, fig. 1.
Luzon, Provinz Laguna, Mount Maquiling. (Belegstiick im
Museum zu Dresden.) Endemische Art.
CALODROMUS MELLYI Guerin.
Calodromus mellyi Guerin, Mag. Zool. (1832) t. 34 c?.
Luzon, Ilocos Norte, Bangui: Manila (Banks).
Die Art is haufig und weit verbreitet. In Indien ist mellyi
haufig und in Bengalen nachgewiesen. Von Burmah bisher
noch nicht bekannt, ist sie in Malakka,
Sumatra und Borneo nicht selten. Von
Java sah ich noch keine Belegstucke.
Calodromus mellyi soil auch auf Ceylon
gefunden sein, ich konnte die Behaup-
tung nicht entkraften, sah aber das Tier
noch nicht von dort.
Die Gattung ist orientalisch. Ver-
wandte Formen finden sich in Sumatra
in der Gattung Allaeodromus Senna
und der aethiopischen Gattung Cormo-
pus Kolbe. Die Gattungen sind durch die tiberbildeten Tarsen
in eine Verwandtschaft zu bringen. In die austro-malayische
Region ist Calodromus nicht vorgedrungen. Habituell besteht
innerhalb der Gattung grosse Einformigkeit.
Genus CYPHAGOGUS Parry
Cyphagogus Parry, Trans. Ent. Soc. London 5 (1849) 182.
CYPHAGOGUS BUCCATUS Kleine.
Cyphagogus buccatus Kleine, Ent. Mitt. 1-4 (1916) 9, figs. 6, 7.
Mindanao, Provinz Lanao, Kolambugan (Boettcher). Sa-
MAR (Baker).
Sicher kommt die Art auch auf anderem Inseln vor, denn sie
ist haufig und weit verbreitet. Ausserdem ist sie leicht erkenn-
bar. In dem mir vorgelegenen Material konnte ich sie nach-
weisen von: Ceylon, Indien, Andamanen, Malakka, Sumatra,
Borneo, und Java. Das Verbreitungsgebiet ist geschlossen, da
buccatus auch von Bengalen nachgewiesen ist.
Fig. 1. Verbreitungskarte der
Gattung Calodromus Guer.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 393
Dresden 1
CYPHAGOGUS EICHHORNI Kirschbaum.
Cyphagogus eichkorni Kirschbaum, Mitt. Zool. Mus.
(1875) 45.
Mindanao, Provinz Lanao, Kolambugan (Baker) ; Provinz
Davao, Davao (C. M. Weber). Negros, Cuernos Mountains
(Baker). Sibuyan (Baker). N.-W.-Panay (Baker).
Wie buccatus ist ,,,
"*3T
ST71
w Werufekrvc
Fig. 2. Verbreitungskarte der Gattung Cyphagogus Parry.
auch eichhorni ver-
breitet und leicht er-
kennbar. Nach Wes-
ten wird Burmah
nicht iiberschritten,
in Indien fehlt sie.
Auf der Malayischen
Halbinsel haufig, ist
sie von Mentawei
und Borneo nachge-
wiesen. Sehr wahr-
scheinlich lebt sie
auch auf Sumatra,
von Java sicher nicht
bekannt. Cyphago-
gus eichhorni ist ferner auf die Molukken tibergegangen, wie
ich selbst feststellen konnte.
CYPHAGOGUS GLADIATOR Kleine.
Cyphagogus gladiator Kleine, Arch. Nat. A 6 87 (1921) 307, figs.
3, 13.
Ich sah die Art von den Philippinen nicht gerade selten,
nahere Fundorte kann ich aber nicht angeben. Die allgemeine
Verbreitung ist der der beiden vorhergehenden Arten ahnlich:
Assam, Malayische Halbinsel, Mentawei, Sumatra und Borneo.
CYPHAGOGUS HUMILIS Kleine.
Cyphagogus hvmilis Kleine, Philip. Journ. Sci. 28 (1925) 590.
Mindanao, Provinz Lanao, Kolambugan (Banks). Ende-
mische Art.
CYPHAGOGUS LONGULUS Senna.
Cyphagogus longulus Senna, Not. Leyd. Mus. 2 (1898) 52.
Mindanao, Provinz Agusan, Cabadbaran (C. M. Weber).
Verbreitung: Siam, Malayische Halbinsel, Java, Ceram, Bat-
jan. Sehr wahrscheinlich ist die Art auch auf Sumatra und
Borneo zu Hause, ich sah aber noch keine Belegstiicke von dort.
394 The Philippine Journal of Science 1931
Der Uebergang von den grossen Sunda-Inseln nach den Philip-
pinen ist ohne Beriihrung von Sumatra und Borneo nich recht
denkbar. Die Verbreitung diirfte der von buccatus und eich-
horni analog sein.
CYPHAGOGUS MODIGLIANII Senna.
Cyphagogus modiglianii Senna, Ann. Mus. Genova (2) 13 (33)
(1893) 258.
Ohne naheren Fundort aus Sammlung Baker.
Eine weit verbreitete, aber zerstreut vorkommende und sel-
tene Art, deren ganzes Verbreitungsgebiet sicher noch nicht
bekannt ist. Senna beschrieb die Art von Sumatra, Insel En-
gano. Ieh sah Stiicke von Pahang, der Fundort uberrascht
nieht. Der Fund von den Philippinen schliesst sich zwanglos
an. Etwas ungereimt erscheint dagagen das Vorkommen in
Nord-Queensland. Es liegt allerdings kein Ausnahmefall vor.
Man kann gleichweite Verbreitung mehrfach feststellen, es
handelt sich dann allerdings um haufige Arten mit grosser
Migration.
CYPHAGOGUS PLANIFRONS Kirschbaum.
Cyphagogus planifrons Kirschbaum, Mitt. Zool. Mus. Dresden 1
(1875) 46.
Mindanao, Provinz Lanao, Iligan (Baker) . Samar (Baker) .
Haufige Art in der weiten Verbreitung von buccatus und eich-
horni: Indien, Assam, Malayische Halbinsel, Borneo, Sumatra,
Java. Das Verbreitungsgebiet ist also gut abgeschlossen.
CYPHAGOGUS SILVANUS Senna.
Cyphagogus silvanus Senna, Boll. Soc. Ent. Ital. 35 (1902) 154.
Bisher lag mir nur einmal ein Stuck aus der Boettcher'schen
Ausbeute mit unleserlichem Fundort von Mindanao vor. All-
gemeine Verbreitung gleieh der vorigen Art: Indien nicht sel-
ten, Malakka, Sumatra, Mentawei, Borneo, Java, Buru.
CYPHAGOGUS SIMULATOR Senna.
Cyphagogus simulator Senna, Boll. Soc. Ent. Ital. 34 (1902) 155.
Mindanao (Baker). Naherer Fundort fehlt.
In der Verbreitung sehr wahrscheinlich mit silvanus iiberein-
stimmend, wenn auch die Funde sich nicht so luckenlos anein-
ander reihen: Assam, Malakka, Sumatra, Borneo und Java.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 395
CYPHAGOGUS TABACICOLA Senna.
Cyphagogus tabacicola Senna, Boll. Soc. Ent. Ital. 25 (1893) 294,
T. 2, fig. 1, lb.
Luzon, Provinz Laguna, Mount Maquiling (Baker). Min-
danao, Provinz Lanao, Kolambugan (Banks).
Eine haufige und leicht erkennbare Art in derselben weiten
Verbreitung wie die vorherigen : Indien, Malakka, grosse Sunda-
Inseln, Andamanen.
CYPHAGOGUS WESTWOODI Parry.
Cyphagogus westwoodi Parry, Trans. Ent. Soc. London 5 (1849)
182.
Negros, Cuernos Mountains (Baker) .
Diese gemeine Art ist auf den Philippinen scheinbar nicht
haufig, denn ich sah sie in dem grossen Material, das mir im
Laufe der Jahre vorgelegen hat, nur dies eine mal. Die Ver-
breitung ist noch ausgedehnter als bei bisher besprochenen
Arten. Von Ceylon bis zu den Philippinen ist sie liickenlos
naehgewiesen und es ist die einzige mir bekannt gewordene Cy-
phagogus-Art, die aus Indo-China gameldet ist und die ich selbst
von dort sah. Das Hauptverbreitungsgebiet ist allerdings
Indien einschliesslich Bengalen, von wo sie fast mit jeder
Bestimmungssendung kommt. Da die Hinterbeine ganz eigen-
artig gebildet sind, so ist keine Verwechslung mit anderen Arten
moglich.
CYPHAGOGUS WHITEI Westwood.
Cyphagogus whitei Westwood, Cab. Or. Ent. (1848) T. 15.
Eine ganz unklare Art, die nur von den Philippinen ( ?) be-
kannt sein soil. Ich konnte sie nach dem grossen Material,
das ich im Laufe der Jahre gesehen habe, nicht identifizieren.
Von den 41 bekannten Arten kommen 12 auf den Philippinen
vor. Sieht man von der unsicheren whitei ab, so bleibt nur eine
Art, die, wenigstens bisher, auf den Inseln endemisch ist.
Jedenfalls wird es aber so sein, dass Endemismen uberhaupt
nicht vorhanden sind.
Die Gattung Cyphagogus hat eine grosse Verbreitung. Von
Ceylon bis Samoa lasst sie sich verfolgen. In Indien und auf
den Sunda-Inseln hat sie eine ansehnliche Artenzahl entwickelt.
In ihrem Grundcharakter ist die Gattung orientalisch, 27
Arten sind dahin zu zahlen, 5 kommen in der orientalischen
396 The Philippine Journal of Science 1&31
und und papuanischen Region vor, der Rest ist ostlich. Selbst
in das Palaearktikum ist die Gattung vorgedrungen, bei Bren-
thiden ein seltener Fall.
Die Cyphagogus der Philippinen sind in ihrer Herkunft leicht
zu deuten: sie sind wahrscheinlich alle von Westen her einge-
drungen. Das ist umsomehr anzunehmen, als die Gattung
grosse Aehnlichkeit mit Cormopus hat, diese liegt aber im Zen-
trum des westlichen Verbreitungskomplexes, in Zentralafrika.
Zu Japan, Celebes oder gar Neu-Guinea bestehen keinerlei Be-
ziehunge, von dort aus sind die Philippinen sieher nicht bevol-
kert worden. Die liber Celebes hinausgegangenen sind zum
grossen Teil in eine ganz andere Farbenentwicklung gekommen.
Nur modiglianii bleibt unklar. Es ist indessen zu beachten,
dass sich auch in Indien und auf den Sunda-Inseln, vernehmlich
an den Randern des Verbreitungskomplexes, bunte Arten finden.
Diese sind aber nicht ohne weiteres mit den Australiern und
Papuanern zu vergleichen, zeigen jedoch, dass die Tendenz,
an den Randgebieten bunte Arten auszubilden, auffallend gross
ist. So erklaren sich auch die vielen bunten Arten in Austra-
lien und dem ostlichen Archipel. Zu den weiten Wanderern
aus dem Westen gehort auch modiglianii; er ist, wie die an-
deren Cyphagogus der Philippinen auch, mit dem grossen Strom
gewandert und, wie es scheint, ein seltener Gast auf verlorenen
Posten geblieben.
Genus EPIGOGUS Kleine
Epigogus Kleine, Ent. Blatt. 19 (1923) 159.
EPIGOGUS FLEXIBILIS Kleine.
Epigoguf flexibilis Kleine, Ent. Blatt. 19 (1923) 159, fig. 1.
Mindanao, Provinz Lanao, Kolambugan (Boettcher). Ne-
GROS, Cuernos Mountains (Baker, Schultze). Basilan (Baker).
Endemische Art.
Genus ORTHOPAREIA Kleine
Orthopareia Kleine, Philip. Journ. Sci. 28 (1925) 591.
ORTHOPAREIA IDONEA Kleine.
Orthopareia idonea Kleine, Philip. Journ. Sci. 28 (1925) 592.
Luzon (Weber). Ohne nahere Fundortangabe. Endemische
Art.
Beide Gattungen umfassen nur je eine Art, es sind also,
wenigstens bis jetzt, auch die Gattungen endemisch.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 397
Genus ASAPHEPTERUM Kleine
Asapheptei-um Kleine, Ent. Mitt. 1-4, 5 (1916) 85.
ASAPHEPTERUM FORMOSANUM Kleine.
Asaphepterum formosanum Kleine, Ent. Mitt. 1-4, 5 (1916) 87, T.
1, figs. 13, 35-37.
Luzon, Provinz Laguna, Mount Banahao (Boettcher).
Diese eigenartige Gattung, die nur diese eine Art umfasst,
fand ich zuerst zahlreich in Formosa-Ausbeuten. Sie ist aber
weiter verbreitet. So sah ich Belegstiicke von Borneo und
Java, mit Ausnahme von Formosa aber immer nur einzeln.
Genus OPISTHENOXYS Kleine
Opisthenoxys Kleine, Arch. Nat. A. 10, 87 (1921) 26.
OPISTHENOXYS BOETTCHERI Kleine.
Opisthenoxys boettcheri Kleine, Philip. Journ. Sci. 28 (1925) 593.
Mindanao, Provinz Zamboanga, Port Banga {Boettcher).
Endemische Art.
OPISTHENOXYS OCHRACEUS Kleine.
Opisthenoxys ochraceus Kleine, Arch. Nat. A. 10, 87 (1921) 28.
Mindanao, Provinz Zamboanga, Zamboanga, Port Banga
(Boettcher) : Provinz Surigao, Surigao
(Boettcher) . Negros, Cuernos Mountains
(Baker), Fabrica (Schultze). Basilan
(Baker) .
Die Art ist haufig und recht weit ver-
breitet : Malayische Halbinsel, Sumatra, Bor-
neo, Java. Die Verbreitung bewegt sich also fig. 3. verbreitunskarte
auf derselben Linie wie die der meisten der Ga"un* oputhe-
noxys Kleine.
Cyphagogus.
Die Gattung umfasst 4 Arten ; die hier nicht genannten kom-
men nur in Indien vor.
Genus PSEUDOCYPHAGOGUS Desbr.
Pseudocyphagogus Desbr., Journ. Asiat. Soc. Beng. 2, 59 (1890) 221.
PSEUDOCYPHAGOGUS SQUAMIFER Desbr.
Pseudocyphagogus squamifer Desbr., Journ. Asiat. Soc. Beng. 2, 59
(1890) 222.
Negros, Cuernos Mountains (Baker). Samar (Baker).
Diese einzige Art der Gattung ist von den Andamanen bes-
chrieben worden und kommt dasalbst auch sehr haufig vor. Die
398
The Philippine Journal of Science
1931
Fig. 4. Verbreitungskarte der
Gattung Mesoderes Senna.
Verbreitung ist aber sehr viel grosser und bewegt sich auf der
Lime der meisten orientalischen Zuwanderer. Mir lag Material
vor von: Assam, Malakka, Sumatra, Borneo.
Genus MESODERES Senna
Mesoderes Senna, Not. Leyd. Mus. 20 (1898) 65.
MESODERES FESSUS Kleine.
Mesoderes fessus Kleine, Ent. Blatt. 19 (1923) 160.
Negros, Cueraos Mountains (Baker.) Endemische Art.
Von den 8 bekannten Arten leben in Indien 2, Malakka 2,
Buru 1, zwei kommen von Malakka bis
Neu-Guinea vor.
Ohne Zweif el ist die Gattung in ihrem
Grundcharakter orientalisch. Der Zug
gegen Osten ist gut zu verfolgen. Die
beiden nicht endemischen Arten mit
grosser Migration lassen deutlich er-
kennen, dass kein sprunghaftes Vord-
ringen stattgefunden hat, denn beide
sind auf alien grossen Sunda-Inseln nachzuweisen. Vielleicht
finden sich aueh auf den Molukken noch einige Arten an.
Genus ATOPOMORPHUS Kleine
Atoporrwrphus Kleine, Philip. Journ. Sci. 28 (1925) 593.
ATOPOMORPHUS SCHULTZEI Kleine.
Atopomorphus schnitzel Kleine, Philip. Journ. Sci. 28 (1925) 594,
t. 1, figs. 1-3.
Negros, Fabrica (Schultze).
N'ur diese eine, auf den Philippinen endemische, Art ist be-
kannt.
Genus ETEROZEMUS Senna
Eterozemus Senna, Boll. Soc. Ent. Ital. 34 (1902') 160.
ETEROZEMUS LAETUS Senna.
Eterozemus laetus Senna, Ann. Mus. Genova (2) 12 (32) (1892)
441.
Luzon, Provinz Nueva Vizcaya, Imugan (Baker).
Weitere sichere Fundorte sind bekannt von Burmah, Sumatra,
Java. Sicher sind die dazwischen liegenden Gebiete auch be-
wohnt, mir lagen aber von der, nicht gerade haufigen, Art keine
Belegstiicke vor.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 399
ETEROZEMUS PUBENS Senna.
Eterozemm pubens Senna, Ann. Mus. Genova (2) 12 (32) (1892)
439.
Luzon, Provinz Laguna, Mount Maquiling (Baker).
Mir lagen Stiicke vor von: Burmah, Perak, Formosa, Java.
Die Verbreitung beider Arten wird wahrscheinlich sehr gleich-
massig sein. Ueber das Verbreitungszentrum der Gattung
lasst sich niehts sicheres sagen, da beide Arten immer nur ein-
zeln gefunden werden. Nur diese beiden Arten sind bekannt.
Genus DICTYOTOPTERUS Kleine
Dictyotopterus Kleine, Ent. Mitt. 1-4, 5 (1916) 75.
DICTYOTOPTERUS PHILIPPINENSIS Kleine.
Dictyotopterus philippinensis Kleine, Arch. Nat. A. 10, 87 (1921)
25.
Philippinen ohne nahere Fundortangabe.
DICTYOTOPTERUS PULCHERRIMUS Kleine.
Dictyotopterus pidcherrimus Kleine, Arch. Nat. A. 10, 87 (1921) 26.
Luzon, Provinz Laguna, Mount Maquiling (Baker) . Ausser-
dem sah ich mehere Belegstucke ohne naheren Fundort.
Es sind drei Arten bekannt. Ausser den beiden genannten
lebt eine Art auf Formosa und den Andamanen. Dictyoto-
pterus entfernt sich also von der grossen Strasse, auf der wir
schon die meisten Zuwanderer kommen sahen, nicht.
Alle auf den Philippinen gefundenen Calodromini sind rein
orientalischen Charakters und mussen als Zuwanderer angesehen
werden. Zwar sind einige, artenarme, Gattungen nur auf den
Philippinen als endemisch festgesteket, aber das will wenig
besagen, da die Abstammung von Orientalen sicher ist.
STEREODERMINI
Genus JONTHOCERUS Lacordaire
Jonthocerus Lacordaire, Gen. Col. 7 (1866) 415.
JONTHOCERUS ASIATICUS Kleine.
Jonthocerus asiaticus Kleine, Arch. Nat. A. S, 85 (1919) 47, figs.
12, 13.
Luzon (ohne nahere Angabe). Mindanao, Provinz Davao
Davao: Provinz Agusan, Butuan (Baker). Palawan, Puerto
Princesa (Sammler unbekannt).
Die Art f and ich f erner aus Material von : Formosa, Sumatra,
Borneo. Ich sah einen Jonthocerus aus Ceylon, der vielleicht
400 The Philippine Journal of Science
Tabelle 3. — Verbreittingstabelle der CalodrommL
1931
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Calodromus crinitus Kleine a
Calodromus mellyi Guerin
Cyphagogus eichhorni Kirschbaum —
Cyphagogus humilis Kleine a _ _ -
Cyphagogus longulus Senna
Cyphagogus modiglianii Senna
Cyphagogus planifrons Kirschbaum . _
Cyphagogus silvanus Senna
Cyphagogus simulator Senna
Cyphagogus tabacicola Senna
Cyphagogus westwoodi Parry
Cyphagogus whitei Westwood •
Epigogus flexibilis Kleine »__- -
Asaphepierum formosanum Kleine
Opisthenoxys boettcheri Kleine a
Opisthenoxys ockraceus Kleine
Pseudocyphagogus squamifer Desbr —
Mesoderes fessus Kleine »
Atopomorphus schultzei Kleine a
Eterozemus Isetus Senna
Eterozemus pubens Senna .
Dictyotopterus philippinensis Kleine a_
Dictyotopterus pulcherrimus Kleine a_ .
a Endemisch.
hierher gehoren konnte. Ich bezweifle aber die Zugehorigkeit,
da nicht einmal von der Malayischen Halbinsel ein Belegstuch
vorlag. Das iibrige Verbreitungsgebiet ist gut abgeschlossen.
Der Uebergang von Borneo nach den Philippinen iiber Palawan
ist interessant. Ueber diese Briicke sind sicher sehr viele Arten
gewandert.
JONTHOCERUS BICOLOR K. M. Heller.
Jonthocerus bicolor K. M. Heller, Deutsche Ent. Zeit. (1916) 297.
Luzon, Provinz Laguna, Mount Banahao (Baker).
Endemische Art. Die Art ist dadurch wiehtig, dass sie die
Ausfarbung der Neu-Guinea-Tiere besitzt. Das kommt auf den
Philippinen ofter vor, in dieser Gattung ist es aber die einzige
Art. Von Neu-Guinea selbst ist kein Jonthocerus bekannt.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 401
JONTHOCERUS LATICOSTATIS Kleine.
Jonthocerus laticostatis Kleine, Arch. Nat. A 8, 85 (1919) 38, figs.
4, 5.
Mindanao, Lanao, Iligan, Kolambugan (Baker).
Ich sah die Art ferner von Formosa, Sumatra, Borneo. Es
ist sehr wahrscheinlich, dass sie westlich bis zur Malayischen
Halbinsel zu finden ist. Die Verbreitung durfte sich mit asia-
ticus ziemlich decken.
Genova (2) 19 (39)
JONTHOCERUS MODIGLIANII Senna.
Jonthocerus modiglianii Senna, Ann. Mus.
(1898) 228.
Mindanao, Provinz Agusan, Butuan (Baker).
Weitere Verbreitung: Andamanen, Sumatra, Mentawei.
Die Gattung umfasst 18 Arten von denen 14 orientalisch sind.
Das Verbreitungszen-
trum liegt auf den I 'Vp^^t u t^kl — J^I/f^l^f
grossen Sunda-Inseln ; * ^* sJCJTv ^} -^_i;*H
von hier aus strahlt
die Gattung iiber For-
mosa bis Japan aus.
Aus Indien habe ich
nur einen recht sehwa-
chen Besatz gesehen,
aber, und das ist wich-
tig, die Verbreitung
ist bis Ceylon, wo noch
eine endemische Art
vorkommt, nicht un-
terbrochen. Das grosse Areal das die Gattung bewohnt ist
daran zu erkepnen, dass sowohl in Afrika wie in Australien je
zwei Arten leben. Der Gattungstyp ist sehr einheitlich.
Genus STEREODERMUS Lacordaire
Stereodermus Lacordaire, Gen. Col. 7 (1866) 419.
STEREODERMUS FLAVOTIBIALIS Kleine.
Stereodermus flavotibialis Kleine, Arch Nat. A. 10, 87 (1921) 28.
Luzon, Provinz Laguna, Mount Maquiling: Provinz Tayabas,
Malinao (Baker).
Endemische Art. Die Gattung ist in der Hauptmasse ihrer
Arten neotropisch, 18 von 25 leben hauptsachlich in Zentral-
Amerika, eine geht bis Siidbrasilien, 6 sind von Senna von den
263774 7
Fig. 5.
Verbreitungskarte der Gattung Jonthocerus
Lacord.
402
The Philippine Journal of Science
1931
Fig, 6. Verbreitungskarte der Gattung Stereodermus Lacord.
Sunda-Inseln beschrieben und ein ist auf den Philippinen en-
demisch. Die Zugehorigkeit aller Arten zur Gattung erscheint
mir hinreichend gesichert. Die Stereodermini sind in alien
grossen Gattungen auffallig weit verbreitet (cfr. Cerobates) .
Genus CEROBATES Schoenherr
Cerobates Schoenherr, Gen. Cure. 5 (1840) 487.
CEROBATES ADUSTUS Senna.
Cerobates adtwtits Senna, Not. Leyd. Mus. 16 (1894) 184.
Mindanao, Provinz Lanao, Iligan {Baker) .
Eine Art von ausserordentlich weiter Verbreitung: Ceylon,
Assam, Malayische Halbinsel, Sumatra, Borneo, Java, Nias,
Bali, Neu-Guinea, Fiji-Inseln. Die Art steht in der Verbreitung
nicht allein da (cfr. sexsulcattts und tristriatus) .
CEROBATES iEQUALIS Kleine.
Cerobates mqualis Kleine, Arch. Nat. A. 3, 87 (1922) 203.
Luzon, Provinz Laguna, Mount Banahao (Boettcher) , Paete
(Boettcher) : Provinz Bataan, Lamao (Boettcher) . Mindanao,
Provinz Lanao, Mumungan (Boettcher) : Provinz Zamboanga,
Port Banga (Boettcher). Samar, Catbalogan (Boettcher) .
Auch diese Art hat eine sehr weite Verbreitung: Ceylon, In-
dien, Indo-China, Malayische Halbinsel, Sumatra, Borneo, Java,
Andamanen, Nicobaren, Ternate. Gegen Osten ist sequalis
nicht so weit vorgedrungen wie adustus.
CEROBATES ANGUSTIPENNIS Senna.
Cerobates angustipennis Senna, Not. Leyd. Mus. 16 (1894) 182.
Luzon, Provinz Laguna, Mount Banahao (Boettcher).
Bisher nur von Java gemeldet. Sehr wahrscheinlich ist die
Art aber, wie die meisten, viel weiter verbreitet und nur noch
nicht aufgefunden worden.
46,s Kleine: Die Brenthiden der Philippinen-Inseln 403
CEROBATES CLINATUS Kleine.
Cerobates clinatus Kleine, Treubia 3-4, 3 (1923) 405.
Mindanao, Provinz Zamboanga, Port Banga (Boettcher) .
Das bei angustvpennis Gesagte gilt auch hier.
CEROBATES COSTATUS Kleine.
Cerobates costatus Kleine, Philip. Journ. Sci. 20 (1922) 153, t. 1,
fig. 2.
Mindanao, Provinz Surigao, Surigao (Baker). Endemische
Art.
CEROBATES FORMOSANUS von Schonfeldt.
Cerobates formosanus von Schonfeldt, Deutsche Ent. Nat. Bibl.
No. 24 2 (1911) 190.
Luzon, Provinz Laguna, Mount Banahao, Los Bafios (Boett-
cher) : Subprovinz Kalinga, Balbalasan (Boettcher). Minda-
nao, Provinz Lanao, Mumungan (Baker). Negros, Cuernos
Mountains (Baker).
Von Schonfeldt hat die Art aus Formosa-Material beschrie-
ben. Auf den Fhilippinen 1st sie aber wenigstens ebenso stark
vertreten wie auf Formosa selbst. Sonst sah ich die Art nicht,
sie schein also auf diesen, verhaltnismassig kleinen, Verbreitung-
skreis beschrankt zu sein.
CEROBATES GROUVELLEI Senna.
Cerobates grouvellei Senna, Boll. Soc. Ent. Ital. 3, 15 (1893) 307,
t. 2, fig. 6.
Mindanao, Provinz Zamboanga, Dapitan (Baker).
Die Art ist sehr weit verbreitet, wenn sich auch kein liicken-
loser Zusammenhang nachweisen lasst. Ich sah Material von:
Sumatra, Borneo, Bali, Queensland.
CEROBATES SEXSULCATUS Motschulsky.
Cerobates sexsulcatus Motschulsky, Et. Ent. 7 (1858) 95.
Luzon, Provinz Laguna, Los Bafios, Paete (Boettcher) . Min-
danao, Provinz Zamboanga, Port Banga (Boettcher) : Provinz
Surigao, Surigao, Dapa (Boettcher) : Provinz Bukidnon, Tang-
kulan (Baker) : Provinz Lanao, Kolambugan (Baker) : Mas-
bate, Aroroy, Cabugao (Boettcher) . Basilan (Baker) .
Eine der gemeinsten und weitverbreitetsten Arten, die genau
erkennen lasst, wie man sich das Wohngebiet der Gattung, so-
weit sie nicht aethiopisch ist, zu denken hat. Es lagen mir
Belegstiicke vor von: Ceylon, Indien, Malayische Halbinsel, Su-
matra, Borneo, Java, Sumbawa, Insel Batu, Andamanen, Cochin-
China, Celebes, Molukken, Neu-Guinea, Queensland.
404
The Philippine Journal of Science
1931
CEROBATES SUMATRANUS Senna.
Cerobates sumatranus Senna, Boll. Soc. Ent. Ital. 3, 25 (1893) 306,
t. 3, fig. 1.
Luzon, Provinz Laguna, Mount Banahao (Boettcher) . Min-
danao, Provinz Zamboanga, Port Banga (Boettcher) : Provinz
Lanao, Mumungan (Boettcher).
Verbreitung der vorigen Art ahnlich: Ceylon, Indien, Mala-
yische Halbinsel, Indo-China, Sumatra, Borneo, Java, Mentawei,
Timor, Formosa, Celebes. Da bereits Celebes und die kleineoi
Sunda-Inseln erreieht sind ist zu erwarten, dass sumatranus noch
weiter ostlich gefunden wird.
CEROBATES TRISTRIATUS Fabricius.
Cerobates tristriatus Fabricius, Syst. El. 2 (1801) 554.
LUZON, Provinz Ilocos Norte, Bangui (Banks) : Provinz Lagu-
na, Mount Banahao, Mount Maquiling (Baker), Magdalena
(Schtdtze), Paete (Boettcher) : Provinz Nueva Vizcaya, Imugan
(Boettcher) : Provinz Bataan, Lamao (Boettcher) , Cabugao
(Boettcher) ; Malinao (Baker) . Mindanao, Provinz Agusan,
Cabadbaran (Weber) : Provinz Lanao, Mumungan (Boettcher) :
Provinz Zamboanga, Port Banga (Boettcher). Siargao, Dapa
(Boettcher). Basilan (Boettcher). Samar (Baker).
Verbreitung also gleich sexsulcatus.
Von den 37 Arten dieser Gattung gehoren 15 der aethiopischen
Kegion an und scheiden ganz aus, 2 sind als australisch anzu-
sehen, 20 sind orientalisch, ganz gleich, wie weit die Verbrei-
tung gegen Osten stattgefunden hat. Der Habitus ist so
einheitlich, dass Zweifel iiber die Zugehorigkeit zur Gattung
ausgeschlossen sind. Die Stereodermini sind ganz allgemein
durch grosse Migration ausgezeichnet. Wenigstens gilt das fur
die artenreichen Gattungen.
Fig. 7. Verbreitungskarte der Gattung Cerobates Schoenherr.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 405
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406
The Philippine Journal of Science
1931
TRACHELIZINI
Genus HOMOPHYLUS Kleine
Homophylus Kleine, Zool. Meded. Leid. 5 (192t)) 244.
HOMOPHYLUS MINDANENSIS Kleine.
Homophylus mmdanensis Kleine, Philip. Journ. Sci. 28 (1925) 595.
Mindanao, Surigao, Surigao (Boettcher). Endemische Art.
Zwei weitere Arten sind von Java bekannt.
Fig. 8. Verbreitungskarte der
Gattung Metatrachelizus Kin.
Genus METATRACHELIZUS Kleine
Metatrachelizus Kleine, Arch. Nat. A. 3, 88 (1922) 207.
METATRACHELIZUS CONSTANS Kleine.
Metatrachelizus constans Kleine, Capita Zool. 4, 2 (1926) 20, t. 1,
fig. 27.
Siargao, Dapa (Boettcher).
Ausserdem von Mysol bekannt.
Uber das wirkliche Verbreitungsa-
raeal der Gattung lasst sich nichts Si-
eheres sagen. Von den 7 Arten sind 4
auf den Molukken, 2 in Indien und 1
auf Borneo gefunden worden. Der
starkere Artbestand auf den Molukken
ist immerhin auffallig und lasst das
Verbreitungszentrum nieht erkennen.
Es besteht eine weitlaufige Ver-
wandtschaft mit den Stereodermini, vielleicht hangt die starke
eigung zur Migration damit zusammen.
Genus TRACHELIZUS Schoenherr
Trachelizws Schoenherr, Gen. Cure. 5 (1840) 489.
TRACHELIZUS BISULCATUS Fabricius.
Brentus bisulcatus (Fabricius), Syst. El. 2 (1801) 548.
Luzon, Provinz Laguna, Mount Maquiling, Malinao (Baker) ;
Mount Banahao (Boettcher) : Provinz Bataan, Lamao (Boett-
cher) : Provinz Nueva Vizcaya, Imugan (Boettcher) . Min-
danao, Provinz Zamboanga, Dapitan (Baker) : Provinz Lanao,
Iligan, Kolambugan (Baker) ; Mumungan (Boettcher) : Provinz
Bukidnon, Tangkulan (Baker). Leyte, Burauen (Boettcher).
Basilan (Boettcher). Samar (Baker).
Der gemeinste Brenthide iiberhaupt. Von Ceylon bis zu den
Salomonen, von Japan bis Queensland. Die Gattung umfasst
46, z Kleine: Die Brenthiden der Philippinen-Inseln 407
11 Arten, von denen keine auch nur entfernt so grosse Migra-
tion aufweist, im Gegenteil, die meisten haben einen kleinen
Verbreitungskreis iiber den sie nicht hinausgehen. Dabei fallt
die Einheitlichkeit im Habitus auf, Stticke von Ceylon sehen
genau so aus wie von Australien, den Salomonen oder Japan.
Genus MIOLISPA Pascoe
Miolispa Pascoe, Journ. Ent. 1 (1862) 393.
MIOLISPA BICOLOR Kleine.
Miolispa bicolor Kleine, Stett. Ent. 80 (1919) 316, fig. 54.
Luzon, Provinz Laguna, Mount Banahao, Mount Maquiling
(Baker): Provinz Nueva Vizcaya, Imugan (Weber). Minda-
nao, Provinz Lanao, Mumungan (Boettcher) : Provinz Surigao,
Surigao (Baker). Mindoro, Subaan (Boettcher). Samar
(Baker) . Endemische Art mit Farbung des Neu-Guineatypus.
MIOLISPA CLAVICORNIS Kleine.
Miolispa clavicornis Kleine, Arch. Nat. A. 10, 87 (1921) 30.
Luzon, Provinz Laguna, Mount Banahao (Boettcher) : Pro-
vinz Nueva Vizcaya, Imugan (Boettcher) . Mindanao, Provinz
Agusan, Butuan (Baker) : Provinz Surigao, Surigao (Boett-
cher): Provinz Lanao, Mumungan (Boettcher). Sibuyan
(Baker). Endemische Art.
MIOLISPA CRUCIATA Senna.
Miolispa cru<ciata Senna, Not. Leyd. Mus. 20 (1898) 69.
Mindanao, Provinz Davao, Davao, (Baker) : Provinz Agusan,
Butuan (Baker) : Provinz Lanao, Mumungan (Boettcher) .
Ich sah die Art ferner von Sumatra, Borneo und Formosa.
Sie ist nicht gerade selten; der Verbreitungsbezirk ist aber
nicht sehr gross.
MIOLISPA DISCORS Senna.
Miolispa discors Senna, Ann. Soc. Ent. Belg. 39 (1895) 358.
Mindanao, Provinz Lanao, Iligan, Cotabato (Taylor; Baker).
Negros, Cuernos Mountains (Baker).
Die Verbreitung ist der vorigen Art ahnlich und nur etwas
ausgedehnter : Penang, Borneo, Formosa, Celebes. Sicher ist
sie auch auf Sumatra zu finden, doch hat mir kein Belegstiiek
vorgelegen. Interessant ist der Fund von Celebes, mit dieser
Insel besteht ofter Uebereinstimmung der Arten.
408 The Philippine Journal of Science 1931
MIOLISPA ELONGATA Kleine.
Miolispa elongata Kleine, Stett. Ent. Zeit. 80 (1919) 244, figs. 13-
17.
Luzon, Manila (Sammler unbekannt). Mindanao, Provinz
Surigao, Surigao (Baker) : Provinz Lanao, Mumungan, Kolam-
bugan (Boettcher) : Provinz Zamboanga, Zamboanga (Baker) :
Provinz Bukidnon, Tangkulan (Boettcher). Negros, Cuernos
Mountains (Baker). Basilan (Baker). Panaon (Baker).
Baker hat die Art ferner im nordlichen Borneo gesammelt.
Der Verbreitungsbezirk scheint aber nur begrenzt zu sein und
die Art muss in erster Linie als philippinisch angesprochen
werden. Wenn sie auf Borneo haufiger ware, hatte sie sich
unter dem grossen Material das ich von Kina Balu gesehen
habe wohl schon einmal gefunden, das ist aber nicht der Fall
gewesen.
MIOLISPA EPHIPPIUM Kleine.
Miolispa ephippium Kleine, Stett. Ent. Zeit. 80 (1919) 247, fig*. 18.
Luzon, Provinz Tayabas, Malinao (Baker) .
Endemisehe Art. Mir lagen spater noch ofter Belegstucke
vor, leider ohne naheren Fundort.
MIOLISPA FLAVOLINEATA Kleine.
Miolispa flavolineata Kleine, Stett. Ent. Zeit. 80 (1919) 282, fig.
37.
Luzon, Provinz Laguna, Mount Banahao, Mount Maquiling
(Baker) : Provinz Nueva Vizcaya, Imugan (Boettcher) . Min-
danao, Provinz Lanao, Uigan, Kolambugan (Baker) . Basilan
(Baker). Endemisehe Art.
MIOLISPA FLEXILIS Kleine.
Miolispa flexilis Kleine, Ent. Blatt. 19 (1923) 161.
Samar (Baker). Mindanao, Provinz Agusan, Butuan
(Baker) : Provinz Lanao, Mumungan (Boettcher) : Provinz
Surigao, Surigao (Boettcher). Endemisehe Art.
MIOLISPA FORMOSA Kleine.
Miolispa formosa Kleine, Ent. Blatt. 19 (1923) 160, figs. 2, 3.
Mindanao, Provinz Agusan, Butuan (Baker). Endemisehe
Art.
MIOLISPA FORNICATA Kleine.
Miolispa fornicata Kleine, Ent. Blatt. 19 (1923) 161.
Luzon, Provinz Laguna, Mount Banahao (Boettcher) : Pro-
vinz Nueva Vizcaya, Imugan (Boettcher) . Mindanao, Provinz
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 409
Lanao, Mumungan (Boettcher) ; Provinz Surigao, Surigao
(Boettcher) . Leyte, Burauen (Boettcher) . Endemische Art.
MIOLISPA FRAUDATRIX Kleine.
Miolispa frmidatrix Kleine, Stett. Ent. Zeit. 80 (1919) 249, fig. 19.
Luzon, Provinz Tayabas, Malinao {Baker). Endemische
Art.
MIOLISPA INTERMEDIA Senna.
Miolispa intermedia Senna, Ann. Soc. Ent. Belg. 41 (1897) 239.
Mindanao, Provinz Lanao, Iligan (Baker) : Provinz Surigao,
Surigao (Baker).
Die Art ist nicht haufig aber doch recht weit verbreitet. Mir
lagen Belegstiicke vor von: Borneo, Java, Celebes und den Mo-
lukken (Amboina).
MIOLISPA LINEATA Senna.
Miolispa lineata Senna, Not. Leyd. Mus. 20 (1898) 57.
Mindanao, Provinz Agusan, Butuan (Baker).
Die Verbreitung dieser Art ist sicher nur ganz mangelhaft
bekannt. Von Java sah ich sie sehr haufig und zwar von alien
Teilen der Insel. Sie kommt aueh auf der Malayischen Halbin-
sel vor, woraus zu sehliessen ist, dass sie wenigstens auf Suma-
tra leben muss.
MIOLISPA PASCOEI Kleine.
Miolispa pascoei Kleine, Stett. Ent. Zeit. 80 (1919) 226.
Luzon, Provinz Laguna, Mount Maquiling, Mount Banahao
(Baker) : Provinz Tayabas, Malinao (Baker). Mindanao, Pro-
vinz Agusan, Butuan (Baker) : Provinz Lanao, Iligan (Baker) .
Endemische Art.
MIOLISPA PAUCICOSTATA Kleine.
Miolispa paucicostata Kleine, Stett. Ent. Zeit. 80 (1919) 312, fig. 52.
Luzon, Provinz Laguna, Mount Maquiling (Baker). Ende-
mische, seltene Art.
MIOLISPA PERSIMILIS Kleine.
Miolispa persimilis Kleine, Philip. Journ. Sci. 20 (1922) 154, t. 1,
fig. 3.
Mindanao, Provinz Lanao, Kolambugan (Baker) ; Mumungan
(Boettcher) . Endemische Art.
410
The Philippine Journal of Science
1931
MIOLISPA PULCHELLA Kleine.
Miolispa pulchella Kleine, Arch. Nat. A. 10, 87 (1921) 29.
Luzon, Subprovinz Benguet, Baguio (Baker) : Provinz La-
guna, Mount Maquiling (Baker) : Provinz Nueva Vizcaya, Imu-
gan (Banks, Boettcher).
MIOLISPA ROBUSTA Kleine.
Miolispa robusta Kleine, Stett. Ent. Zeit. 80 (1919) 230, fig. 8.
Luzon, Provinz Laguna, Los Banos, Mount Banahao (Boett-
cher). Catanduanes, Virac (Boettcher). Mindanao, Prov-
inz Davao, Davao, (Baker) : Provinz Surigao, Surigao (Baker) :
Provinz Agusan, Butuan (Baker) : Provinz Lanao, Kolambugan,
Uigan (Baker) ; Mumungan (Boettcher) ; San Miguel (Boett-
cher). Mindoro, Subaan (Boettcher). Siargao, Dapa (Boett-
cher). Polillo (Boettcher). Samar (Baker). Basilan
(Baker).
Die Art ist sicher auf den Philippinen zu Hause. Das Auf-
finden auf Borneo (Sandakan) durch Baker beweist mir, dass
die Verbreitung nur gering ist, sonst waren weitere Funde nach-
gewiesen. (cfr. elongata).
MIOLISPA SIPORABA Senna.
Miolispa siporaba Senna, Ann. Mus. Genova (2) 19 (39) (1898) 233.
Mindanao, Surigao, Surigao (Baker).
Eine der verbreitetsten Arten, die sich schon deutlich auf
dem mehrfach skizzierten Wege von Westen nach Osten bewegt.
Ich sah sichere Belegstiicke von Malakka, Sumatra, Mentawei,
Borneo, Java und den Molukken.
MIOLISPA UNICOLOR Kleine.
Miolispa unicolor Kleine, Stett. Ent. Zeit. 80 (1919) 314, fig. 53.
Luzon, Provinz Laguna, Mount
Banahao (Baker) : Provinz Nueva
Vizcaya, Imugan (Boettcher).
Endemische Art.
Die Gattung Miolispa ist die ar-
tenreichste der ganzen Familie.
Von den bekannten 65 Arten kom-
men 20 auf den Philippinen vor
und davon 13 als Endemismen.
Von den restlichen 8 mussen we-
nigstens noch 2 (elongata und ro-
busta) als Philippinentiere angesprochen werden. Nur einige
V^v
-Y^
&1
r—
Vs
* **JW\ J
«0
.i
*
7^
It
**V
■■{
— *
J
Fig. 9.
Verbreitungskarte der Gattung
Miolispa Pascoe.
46,8 Kleine: Die Brenthiden der Philippinen-Inseln 411
lassen den bekannten Weg aus dem ostlichen Verbreitungszen-
trum erkennen.
Das Verbreitungsareal der Gattung ist gross. Ceylon und
Indien haben merkwiirdigerweise keine Vertreter. Erst in Bur-
mah linden sich die ersten Spuren, die aber aueh nur als Aus-
laufer anzusehen sind. Das Verbreitungszentrum liegt auf dem
siidlichen Teil der Malayischen Halbinsel und den grossen Sun-
da-Inseln. Von hier aus ist starke Abwanderung sowohl nach
den Philippinen wie nach Celebes und ins austro-malayische
und australische Gebiet festzustellen. Bemerkenswert ist fur
die Gattung die Tatsache, dass sich so zahlreiche Arten mit
kleinem Verbreitungskreis gebildet haben. Neigung zur Varia-
tion ist allgemein gering.
Genus HYPOMIOLISPA Kleine
Hypomioligpa Kleine, Ent. Blatt. 14 (1918) 163.
HYPOMIOLISPA EXARATA Desbr.
Hypomiolispa exarata Desbr., Journ. Asiat. Soc. Beng. 2, Nat. Sc. No.
3 (1890) 223.
Mindanao, Provinz Zamboanga, Zamboanga (Baker) : Pro-
vinz Lanao, Iligan (Baker) : Provinz Surigao, Surigao (Baker).
Samar (Baker). Basilan (Baker).
Ausser auf den Philippinen, Sumatra, Borneo und Java ge-
funden. Auf Sumatra und Java ist die Art sehr haufig, auf
Borneo lasst die Besatzstarke nach und die Philippinen konnen
nur noch als vorgeschobener Post en angesehen werden.
HYPOMIOLISPA HELLERI Kleine.
Hypomiolispa helleri Kleine, Ent. Blatt. 14 (1918) 329.
Mindanao, Provinz Davao, Davao (Baker) : Provinz Surigao,
Surigao (Baker). Basilan (Baker). Endemische Art.
HYPOMIOLISPA NUPTA Senna.
Hypomiolispa nupta Senna, Not. Leyd. Mus. 14 (1892) 171.
Luzon, Provinz Tayabas, Malinao (Baker) . Mindanao, Pro-
vinz Lanao, Kolambugan, Mumungan (Boettcher) : Provinz
Agusan, Butuan (Baker).
Eine der verbreitetsten Arten auf der West-Ost-Strasse sehr
gut nachweisbar: Assam, Malayische Halbinsel, Sumatra, Bor-
neo, Java, Mentawei.
412 The Philippine Journal of Science 1931
HYPOMIOLISPA OCULARIS Kleine.
Hypomiolispa ocularis Kleine, Proc. Hawaiian Ent. Soc. (1) 7 (1927)
(1928) 57, figs. 2, 3.
Luzon, Provinz Laguna, Los Banos (Sammler unbekannt).
Endemische Art.
HYPOMIOLISPA SPONSA Kleine.
Hypomiolispa sponsa Kleine, Ent. Blatt. 14 (1918) 324.
Mindanao, Provinz Surigao, Surigao {Baker) : Provinz La-
nao, Mumungan (Boettcher) . Samar (Baker).
Diese mit nupta verwandte Art hat fast dieselbe Verbreitung :
Malayische Halbinsel, Sumatra, Borneo, Java, Mentawei; die
Westgrenze seheint aber schon in Selangor zu liegen.
HYPOMIOLISPA TOMENTOSA Kleine.
Hypomiolispa tomentosa Kleine, Philip. Journ. Sci. 20 (1922) 156.
Mindanao, Provinz Lanao, Iligan (Baker). Endemische
Art.
HYPOMIOLISPA TRACHELIZOIDES Senna.
Hypomiolispa trachelizoides Senna, Not. Leyd. Mus. 16 (1894) 193.
Mindanao, Provinz Agusan, Butuan (Baker) : Provinz La-
nao, Iligan (Baker) ; Mumungan (Boettcher) . Samar (Baker) .
Verbreitung gleich sponsa, ausserdem noch auf Celebes ge-
funden. Verbreitungszentren sind die grossen Sunda-Inseln
mit Sumatra als Hauptinsel.
Die Gattung umfasst 32 Arten. Mit Miolispa besteht einige,
wenn auch entfernte, Verwandtschaft. Einige Arten waren
friiher bei Miolispa untergebracht. Ganz zu Unreeht. Die
zoogeographischen Verhaltnisse sind in beiden Gattungen total
verschieden. Miolispa ist stark nach Osten, Hypomiolispa nach
Westen orientiert. Nur im Verbreitungszentrum beider Gat-
tungen, den Sunda-Inseln, treffen sie in etwa gleicher Starke
zusammen. In der auf den Philippinen festgestellten Arten-
zahl dokumentiert sich das Gesagte. Wahrend Miolispa noch
mit der ansehnlichen Zahl von 20 Arten vorkommt, hat es Hypo-
miolispa nur auf 7 gebracht, davon 2 Endemismen. Auf den
Philippinen liegt auch die Ostgrenze der Verbreitung, nach Su-
dosten ist die Gattung nicht vorgedrungen, Farbungselemente
die auf Neu-Guinea hinweisen und die sich in der Gattung
Miolispa mehrfach finden, fehlen ganz.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 413
Genus HIGONIUS Lewis
Higonius Lewis, Journ. Linn. Soc. Lond. Zool. 17 (1883) 299.
HIGONIUS CILO Lewis.
Higonius cilo Lewis, Journ. Linn. Soc. Lond. Zool. 17 (1883) 300, t.
12, figs. 9, 10.
Luzon, Subprovinz Kalinga, Balbalasan (Boettcher) . Masba-
te, Aroroy (Boettcher).
Die Verbreitung dieser Art ist eigenartig: Indien, Burmah,
Formosa, Japan. Von den Sunda-Inseln und Malakka, von wo
ieh so viel Material gesehen habe, ist sie mir niemals vorgekom-
men. Andere Arten, zum Beispiel, crux, haben sich auf dieser
Strasse ausgebreitet. Higonius ist rein orientalisch.
Genus MICROTRACHELIZUS Senna
Microtrachelizm Senna, Boll. Soc. Ent. Ital. 25 (1893) 315.
MICROTRACHELIZUS FLUXUS Kleine.
Microtrachelizus fluxus Kleine, Ent. Blatt. 19 (1923) 162.
Negros, Cuernos Mountains (Baker). Endemische Art.
MICROTRACHELIZUS PUBESCENS Senna.
Microtrachelizus pubescens Senna, Boll. Soc. Ent. Ital. 25 (1893)
320, t. 3, fig. 6.
Negros, Cuernos Mountains (Baker).
Weitere Fundorte sind bekannt von: Malakka (Perak) und
Sumatra.
MICROTRACHELIZUS SIAMENSIS Kleine.
Microtrachelizus siamensis Kleine, Journ. Fed. Malay Stat. Mus. 2
and 3, 13 (1926) 165, fig. 3.
Samar, Catbalogan (Boettcher).
Die Art lag mehrfach von der Malayischen Halbinsel vor,
die Verbreitung durfte sich mit der von pubescens decken.
MICROTRACHELIZUS TABACI Senna.
Microtrachelizus tabaci Senna, Boll. Soc. Ent. Ital. 25 (1893) 323,
t. 4, %. 4.
Mindanao, Provinz Zamboanga, Port Banga (Boettcher) .
Weitverbreitete Art. Ich sah Belegstiicke von : Burmah Ma-
lakka, Sumatra und Borneo. Der Autor nennt Neu-Guinea.
Die Gattung umfasst 27 Arten von denen 4 auf den Philip-
pinen nachgewiesen sind. Die als Endemisme bezeichnete Art
414
The Philippine Journal of Science
1931
ist wahrseheinlich auch noch auf den Sunda-Inseln zu finden.
Die allgemeine Verbreitung ist f olgende : 6 Arten sind eethiopisch
und zeigen noch die Herkunft an, 15 sind rein orientalisch, 3
gehoren dem austro-malayischen beziehungsweise dem austra-
lischen Gebiet an und 3 kommen in sehr grosser Migration
iiber meherer Gebiete vor. Die sethiopischen Vertreter der Gat-
tung sind in sich abgesehlossen sind aber habituel mit den an-
derer Regionen absolut tibereinstimmend. Die philippinischen
Arten sind nur Auslaufer des auf den Sunda-Inseln liegenden
Massivs der orientalischen Arten und sind zoogeographiseh
ohne Belang. Von Bedeutung ist der Naehweis der Afrikaner.
Microtrachelizus ist iibrigens nicht die einzige Gattung die noch
in Afrika Vertreter hat. Es sei nur auf Araiorrhynchus ver-
wiesen, die allerdings die Philippinen nieht erreicht hat.
Fig. 10. Verbreitungskarte der Gattung Microtrachelizus Senna.
Genus HOPLOPISTHIUS Senna
Hoplopisthius Senna, Ann. Mus. Genova (2) 12 (33) (1892) 451.
HOPLOPISTHIUS TRICHIMERUS Senna.
Hoplopisthius trichimerus Senna, Ann. Mus. Genova (2) 12 (33)
(1892) 451.
Luzon, Provinz Laguna, Mount Maquiling (Baker). Pala-
wan, Puerto Princesa (Baker) .
Sehr verbreitete Art. Folgende Fundorte sind bekannt: As-
sam, Burmah, Malayische Halbinsel, Sumatra, Borneo, Java,
Mentawei, Bali, Nias, Formosa.
Eine zweite Art ist von Celebes bekannt. Der Uebergang
von den Sunda-Inseln uber Palawan nach den Philippinen ist
wichtig.
46,3
Kleine: Die Brenthiden der Philippinen-Inseln 415
Fig. 11. Verbreitungskarte der Gattung Cordus Schoenh.
AMORPHOCEPHALINI
Genus CORDUS Schoenherr
Cordus Schoenherr, Mant. Insec. Cure. (1847) 10.
CORDUS PEGUANUS Senna.
Cordus peguanus Senna, Ann. Mus. Genova (2) 12 (32) (1892) 463.
Negros, Cuernos Mountains (Baker). Basilan (Baker).
Weitere Verbreitung: Burmah, Malakka, Sumatra.
Die Gattung ist da-
d u r c h interessant, j — j-p£
dass sie in sehr weiter
Verbreitung vorkommt
und dazwischen in
grossen Gebieten wie-
der ganz fehlt. Zehn
Arten sind aethiopiseh,
2 orientalisch, 1 ist
von Neu-Guinea be-
kannt und 7 sind Aus-
tralier. Das schwache
Auftreten im orienta-
lischen Gebiet ist noch
ganz ungeklart. Das Areal von Malakka und den grossen
Sunda-Inseln ist so intensiv durchforscht, dass langst ein Cordus
bekannt geworden ware, wenn einer vorhanden ware. Andere
Amorphocephalini sind doch mehrfach aufgefunden. Diese Gat-
tung muss einen Weg genommen haben, der heute nicht nehr er-
kennbar und in den einzelnen Erdperioden verloren gegangen ist.
Das sich auf den Philippinen nicht einmal eine eigene Art fand,
ist merkwiirdig, zeigt aber, dass einzelne Arten eine recht grosse
Verbreitung haben konnen. Jedenfalls ist der Cordus von den
Philippinen ein sehr interessanter Fall.
Genus LEPTAMORPHOCEPHALUS Kleine
Leptamorphocephalus Kleine, Arch. Nat. A. 12, 82 (1916) (1918)
132.
LEPTAMORPHOCEPHALUS FCEDERATUS Kleine.
Leptamorphocephalus fcederatus Kleine, Ent. Blatt. 19 (1923) 163.
Negros, Cuernos Mountains (Baker). Endemische Art.
Die 9 Arten umfassende Gattung ist rein orientalisch, mit
ihrer Hauptstarke auf der Malayischen Halbinsel und Sumatra ;
foederatus ist ein vorgeschobener Posten.
416
The Philippine Journal of Science
1931
g
•8
05
03
4«, 3 Kleine: Die Brenthiden der Philippinen-Inseln 417
I I
I I
I I
I I
I I I
I I I
I +
I I I
! I I
I I I
I +
MM
+ +
I I + +
+ + + +
+ I + +
MM
2 | M |
S -8
I «
.a J s,
1
•8
a o e e «
O ^ ^ v vj
&, p, e, a, g, ,s
•3 .3 -S -8 -S o ^ ^
•2 -3 -2 -2 »2
§ 3
263774 8
418 The Philippine Journal of Science mi
Genus PARAMORPHOCEPHALUS Kleine
Paramorphocephahis Kleine, Zool. Meded. Leid. 4, 5 (1920) 236.
PARAMORPHOCEPHALUS SETOSUS Kleine.
Paramorphocephahis setosus Kleine, Philip. Journ. Sci. 28 (1925)
597, taf. 1, fig. 4.
Samar (Baker.) Endemische Art.
Die Amorphocephalini sind Myrmecophile, ihre Verbreitung
ist also immer mehr oder weniger von der ihrer Wirtstiere
abhangig. Letztere sind leider nur erst ganz wenig bekannt.
Auf den Philippinen ist die Tribus sicher nur ganz schwach
vertreten. Jahrelang haben mir iiberhaupt keine Vertreter vor-
gelegen, erst spater konnte ich den Nachweis erbringen, dass
sich auch auf diesen vorgesehobenen Posten myrmecophile Bren-
thiden finden. Es handelt sich in jedem Fall urn Vorposten aus
dem orientalischen Massiv, das sich um die Malayische Halbinsel
und Sumatra konzentriert. Nach den Randern des Gebietes
lasst die Artstarke nach. Etwa die Halfte aller Amorphoce-
phalini leben in Afrika. Keine Tribus der ganzen Familie hat
iibrigens eine derartige West-Ost-Ausdehnung wie die Amor-
phocephalini : Bucht von Guinea bis Tahiti !
ARRHENODINI
Genus AGRIORRHYNCHUS Power
Agriorrhynchus Power, Pet. Nouv. Ent. 2 (1878) 241.
AGRIORRHYNCHUS IGNARIUS Kleine.
Agriorrhynchus ignarius Kleine, Philip. Journ, Sci. 28 (1925) 598,
t. 1, figs. 5-7.
Luzon, Provinz Laguna, Los Baiios (Banks). Endemische
Art.
Vier Arten sind bekannt, alle sind Orientalen die westlich
nicht liber Burmah hinausgehen. Verbreitungszentrum sind
die grossen Sunda-Inseln.
Genus EUPEITHES Senna
Eupeithes Senna, Ann. Mus. Genova (2) 19 (39) (1898) 381.
EUPEITHES DOMINATOR Kleine.
Eupeithes dommator Kleine, Ent. Blatt. 17 (1921) 125, fig. 4.
Mindanao, Provinz Surigao, Surigao (Baker). Samar,
Wright (McGregor) . Endemische Art.
Vier Arten sind bekannt. Ueber die Verbreitung gilt das bei
Agriorrhynchus Gesagte.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 419
Genus PROPHTHALMUS Lacordaire
Prophthalmus Lacordaire, Gen. Col. 7 (1866) 427.
PROPHTHALMUS LONGIROSTRIS Gyllenhal.
Prophthalmus longirostris Gyllenhal, Schoenh. Gen. Cure. 1 (1833)
323.
Luzon, Provinz Laguna, Mount Banahao (Baker).
Auf den Sunda-Inseln gemein, ferner von der Malayischen
Halbinsel und Celebes bekannt. Es ist in der Gattung die Art
mit grosster Migration.
PROPHTHALMUS TRICOLOR Power.
Prophthalmus tricolor Power, Ann. Soc. Ent. Fr. (5) 8 (1878) 38.
Luzon, Provinz Camarines Sur, Mount Isarog (Boettcher) ;
Provinz Laguna, Mount Maquiling (Baker), Mount Banahao,
Los Banos, Paete (Boettcher) : Provinz Ilocos Norte, Bangui
(Boettcher) : Provinz Pampanga, Arayat (Boettcher). Catan-
duanes, Virac (Boettcher). Mindanao, Provinz Surigao, Suri-
gao (Boettcher): Provinz Lanao, Mumungan (Boettcher).
Siargao, Dapa and Cabuntog (Boettcher). Negros, Cuernos
Mountains (Baker). Samar (Bak^). Leyte (Boettcher).
Sibuyan (Baker).
Die Art ist auf den Philippinen eine der haufigsten Brenthi-
den, die sieher auf alien Inseln zu finden ist. Die Art kommt
ferner auf Celebes und auf den Molukken vor (Ceram, Buru,
Amboina) . Es haben sieh Rassen gebildet, die aber keine geo-
graphischen Sehliisse zulassen.
Von den bekannten 17 Arten ist nur eine austromalayisch,
alle anderen sind orientalisch.
Genus BARYRRHYNCHUS Lacordaire
Baryrrhynchus LACORDAIRE, Gen. Col. 7 (1866) 428.
BARYRRHYNCHUS SCHROEDERI Kleine.
Baryrrhynchus schroederi Kleine, Stett. Ent. Zeit. (1914) 172.
Luzon, Provinz Laguna, Mount
Maquiling, Los Banos (Baker) ;
Mount Banahao (Boettcher) : Sub-
provinz K a 1 i n g a , Balbalasan
(Boettcher). Mindanao, Provinz
Agusan, Agusan (Weber) : Prov-
inz Surigao, Surigao (Boettcher) :
Provinz Lanao, Mumungan, Ko-
lambugan (Boettcher). NEGROS,
Cuernos Mountains (Baker)
(Baker) .
V^.
s^tf^
)
> vijRT Jfe/**
.1
•;;:
\ .«
Fig. 12. Verbreitnngskarte der Gat-
tung Baryrrhynchus Lacord.
Sibuyan (Baker) . Samar
420
The Philippine Journal of Science
1931
Baryrrhynchus schroederi ist eine der interessantesten Arten
in der Gattung, denn sie hat eine Verbreitung wie keine andere.
Ich sah sie auf folgender Linie: Siam-Philippinen-Celebes-Mo-
lukken-Neu-Guinea-Neu-Pommern. Sie umgeht also das orien-
talische Gebiet, denn der Vorstoss gegen Siam hat ganz bestim-
mut nicht uber die Sunda-Inseln stattgefunden, sondern uber
Indo-China, wo die Art sehr wahrscheinlich noch aufgefunden
wird. Sie ist auch in der Ausfarbung ganz appart. Von den
18 Arten sind 11 orientalisch, 7 gehoren dem austro-malayischen
beziehungsweise australischen Gebiet an. Die Orientalen haben
keine Fiihlung mit den ostliehen Arten.
Genus EUPSALIS Lacordaire
Eupsalis Lacordaire, Gen. Col. 7 (1866) 430.
EUPSALIS KLEINEI K. M. Heller.
Eupsalis kleinei K. M. Heller, Philip. Journ. Sci. 19 (1921) 624, t.
3, figs. 13, 14.
Mindanao, Provinz Davao, Davao (im Museum Dresden,
wahrscheinlich von Baker gesammelt) . Endemische Art.
Die Gattung ist kein einheitlicher Typ. Die auf den Moluk-
ken, auf Neu-Guinea und Australien lebenden Arten sind der
Untergattung Schizoeupsalis zuzuweisen. Hierher gehort auch
kleinei Heller. Die Beeinflussung durch ostlicher Elemente ist
ganz sicher, die beiden orientalischen Arten, von denen eine nur
in Indien vorkommt, sind ohne Einfluss geblieben. Die Haupt-
masse, 18 Arten, sind sethiopisch mit Ausstrahlung ins medi-
terrane Gebiet. Die orientalen gehoren habituel noch dem afri-
kanischen Artmassiv an, die von den Philippine siidlich und
Fig. 13. Verbreitungrskarte der Gattung EupsaMa Lacord.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 421
siidostlich vorkommenden Arten bilden einen eigenen Verwandt-
schaftskreis. Sie sind am besten von Eupsalis zu trennen.
Genus CiENORYCHODES Kleine
Cssnorychodes Kleine, Arch. Nat. A. 9, 86 (1920) 87.
CiENORYCHODES SERRIROSTRIS Fabricius.
Casnorychodes serrirostris Fabricius, Syst. El. 2 (1801) 553.
Luzon, Provinz Laguna, Mount Maquiling (Baker) . Masba-
te, Aroroy (Boettcher). Mindanao, Provinz Agusan, Agusan
River (Weber) : Provinz Lanao, Iligan, Mumungan, Kolambu-
gan, (Boettcher) : Provinz Cotabato, Cotabato (Taylor, Baker).
Leyte (Boettcher). Siargao, Dapa (Boettcher). Samar
(Baker). Basilan (Baker).
Weitverbreitete gemeine Art, die sich durch ihr grosses Mi-
grationsvermogen bis Celebes vorgeschoben hat. Verbreitungs-
gebiet: Malakka, Sumatra, Borneo, Java, Batu, Bali, Indo-China,
Formosa, Obir.
CJENORYCHODES SPLENDENS Kirsch.
Casnorychodes spleTidens Kirsch., Mitt. Zool. Mus. Dres. 1 (1875) 50
nota.
Luzon, Provinz Laguna, Mount
Banahao (Boettcher) : Provinz
Camarines Sur, Mount Isarog
(Boettcher) . Masbate, Aroroy
(Boettcher) . Negros, Cuernos
Mountains (Baker). Sibuyan
(Baker). Endemische Art.
Zwolf Arten sind bekannt, davon
sind 6 orientalisch, 6 austromalay-
isch. Die Arten haben allgemein
wenig Neigung zu Migration, nur serrirostris ist Weiter ver-
breitet.
Genus PSEUDORYCHODES Senna
P8eudorychodes Senna, Ann. Soc. Ent. Belg. 38
(1894) 375.
PSEUDORYCHODES PRiECLARUS Kleine.
Pseudorychodes prseclarus Kleine, Philip. Journ.
Sci. 28 (1925) 600, t. 1, fig. 8.
Mindanao, Provinz Surigao (Boettcher) .
Endemische Art.
Von den 13 Arten sind 11 orientalisch, 1 ist
von Celebes, 1 von Japan. Bei keiner Art ist
s~
?f
r ■
•
V
si
'*/**
^x
» tj .'
•;t
"~s
• v •
Fio.
14. Verbreitungskarte der
tung Caencrychodes Kin.
Gat-
Fig. 15. Verbrei-
tungskarte der
Gattung Peevr
dorychodes Sen-
na.
grosse Migration erkennbar.
422
The Philippine Journal of Science
1931
Genus AMPHICORDUS K. M. Heller
Amphicordus K. M. Heller, Philip. Journ. Sci. § D 8 (1913) 151.
AMPHICORDUS IMPROPORTIONALIS K. M. Heller.
Amphicordus improportionalis K. M. Heller, Philip. Journ. Sci. § D
8 (1913) 152, fig. 7.
Mindanao ohne naheren Fundort (Museum Dresden) ; Pro-
vinz Lanao, Kolambugan (Banks) . Nur diese erne, endemische,
Art ist bekannt.
Tabelle 6. — Verbreitungstabelle der Arrhenodini,
a
o
6
-■d
a
i
o
g
*-,
o
"ctf
2
I | | + ! I 1 + ||| Sumatra.
o
<U
c
o
>
to
O
c
o
"o
c
O
55
Agriorrhynchns ignarius Kleine a
Eupeithes dominator Kleine a
Prophthalmu8 longirostris Gy-
Uenhal
+
+
+
+
+
+
+
+
+
+
+
+
+
+
Prophthalmus tricolor Power
Baryrrhynchus schroederi Kleine
Eupsalis kleinei Heller a
Cxnorychodes serrirostris Fa-
bricius._
Csenorychodes splendens Kirscha
Pseudorychodes prmclarus
Kleine 8
A mphicordus improportionalis
Heller *
a Endemisch.
Von den 195 bekannten Arrhenodini sind 90 orientalisch.
Der Bestand auf den Philippinen mit nur 10 Arten ist also sehr
gering. Schuld mag daran der Umstand mit sein, das in der
Tribus so wenig Neigung zu Migration besteht.
BELOPHERINI
Genus YPSELOGONIA Kleine
Ypselogonia Kleine, Philip. Journ. Sci. 20 (1922) 157.
YPSELOGONIA PEREGRINA Kleine.
Ypselogonia peregrina Kleine, Philip. Journ. Sci. 20 (1922) 158, t.
1, fig. 1.
Mindanao, Provinz Zamboanga, Dapitan (Baker). Ende-
mische Art.
Eine zweite Art kommt in Formosa und Borneo vor.
46, '5 Kleine: Die Brenthiden der Philippinen-Inseln 423
Genus HETEROBLYSMIA Kleine
Heteroblysmia Kleine, Ent. Blatt. 13 (1917) 285.
HETEROBLYSMIA ACCURATA Kleine.
Heteroblysmia accurata Kleine, Arch. Nat. A. 3, 88 (1922) 215.
Mindanao, Provinz Lanao, Kolambugan (Boettcher).
Die Art habe ich mehrfach von Borneo gesehen, wo sicher
das Verbreitungszentrum liegt. Es ware von Interesse fest-
zustellen, ob sich accurata nicht auch auf Palawan findet.
HETEROBLYSMIA ELECTA Kleine.
Heteroblysmia electa Kleine, Ent. Blatt. 19 (1923) 164, figs. 5, 6.
Naherer Fundort fehlt. Endemische Art.
HETEROBLYSMIA FORMIDOLOSA Kleine.
Heteroblysmia formidolosa Kleine, Ent. Blatt. 19 (1923) 165, fig. 7.
NEGROS, Cuernos Mountains (Baker). Endemische Art.
Acht Arten sind bekannt, rein orientalisch und meist von den
Sunda-Inseln.
Genus APOCEMUS Calabresi
Apocemus Calabresi, Boll. Soc. Ent. Ital. 53 (1929) 58.
APOCEMUS IGNOBILIS Kleine.
Apocemus ignobilis Kleine, Philip. Journ. Sci. 28 (1925) 602, t. 1,
fig. 9.
Luzon, Provinz Bataan, Lamao (Carpenter). Endemische
Art. Eine zweite Art ist von Malakka bekannt.
Genus HENARRHENODES K. M. Heller
Henarrhenodes K. M. Heller, Philip. Journ. Sci. § D 8 (1913) 152.
HENARRHENODES MACGREGORI K. M. Heller.
Henarrhenodes macgregori K. M. Heller, Philip. Journ. Sci. § D 8
(1913) 153, fig. 8.
Luzon, Subprovinz Benguet, Irisan River (Baker) : Provinz
Nueva Vizcaya, Imugan (Baker) : Provinz Laguna, Mount Ma-
quiling (Baker). Mindanao, Provinz Lanao, Kolambugan
(Baker). Siargao, Cabuntog (Boettcher). Negros, Cuernos
Mountains (Baker). Polillo (Sammler unbekannt, wahrsche-
inlich Baker).
Es sind noch 2 orientalische Arten bekannt. Die philippi-
nische Art ist dadurch ausgezeichnet, dass sie den Ausfarbungs-
typ der Neu-Guinea-Tiere hat.
424 The Philippine Journal of Science 1931
Genus ECTOCEMUS Pascoe
Ectocemus Pascoe, Journ. Ent. 1 (1862) 388.
ECTOCEMUS BADENI Kirsch.
Ectocemus badeni Kirsch, Ent. Mitt. Mus. Dresden 1 (1875) 48.
Luzon, Provinz Laguna, Mount Banahao (Baker). Minda-
nao, Provinz Lanao, Kolambugan (Banks): Surigao (Baker).
Die Gattung umfasst 5 Arten, 3 sind orientalisch, 2 austro-
malayisch oder australisch.
Genus ANEPSIOTES Kleine
AnepsMes Kleine, Ent. Mitt. 10-12, 6 (1917) 318.
ANEPSIOTES fcUZONICUS Calabresi.
Ampsiotes luzonicus Calabresi, Boll. Soc. Ent. Ital. 51 (1919) 66,
t. 2, fig. 3.
Luzon (Coll. Senna). Endemische Art.
ANEPSIOTES NITIDICOLLIS Calabresi.
Anepeiotes nitidicollis Calabresi, Boll. Soc. Ent. Ital. 51 (1919)
(1929) 69, t. 2, fig. 4.
Manila (Coll. Senna). Endemische Art.
Die Gattung umfasst 7 orientalische Arten mit geringer Mi-
gration.
Tabelle 7. — Verbreitungstabelle der Belopherini.
Borneo.
Celebes.
Ypselogonta peregrina Kleine » «
+
+
Heteroblysmia accurate Kleine
Heteroblysmia electa Kleine * .
Heteroblysmia formidolosa Kleine a
Apocemu8 iguobilis Kleine »
Ectocemus badeni Kirsch
Anepsiotes luzonicus Calabresi »
Anepsiotes nitidicollis Calabresi *
11 Endemisch.
Die Belopherini sind von den bisher behandelten Tribus grund-
eatlich dadurch unterschieden, dass die nicht westlicher, son-
dern siidostlicher Provenienz sind. Die Tribus findet sich cir-
cumpolar auf der ganzen siidlichen Hemisphere mit Ausnahme
von Afrika (wohl aber in Madagaskar) . Der heute noch deut-
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 425
lich erkennbare Wanderzug der in die orientalische Region ge-
kommen ist stammt aus dem Neu-Guinea-Massiv, auf das ich
schon mehrfach hingewiesen habe. Von hieraus hat auch die
Besiedelung der Philippinen stattgefunden, reine Typen Neu-
Guineas finden sich und der Mangel jeder Anlehnung an die
orientalische Region lassen das auch deutlich erkennen. Von
96 bekannten Arten sind 34 in die orientalische Region vorge-
drungen und haben dort zum Teil ganz neue Formen gebildet.
ITHYSTENINI
Genus ACHRIONOTA Pascoe
Achrionota Pascoe, Ann. & Mag. Nat. Hist. (4) 10 (1872) 325.
ACHRIONOTA BILINEATA Pascoe.
Achrionota bilineata Pascoe, Ann. & Mag. Nat. Hist, (4) 10 (1872)
325, t. 15, fig. 4.
Mindanao, Provinz Zamboanga, Dapitan (Baker).
Weiter haben mir folgende Fundorte vorgelegen: Malakka,
Sumatra, Borneo.
ACHRIONOTA SPINIFER Kleine.
Achrionota spinifer Kleine, Arch. Nat. A, 10, 87 (1921) 36, figs. 2,
3.
Mindanao, Provinz Surigao, Surigao (Baker). Siargao,
Dapa (Boettcher). Leyte (Boettcher). Panaon (Boettcher).
Bohol, Bilar (Ramos). Polillo (McGregor). Endemische
Art. Eine dritte Art kommt endemisch auf Celebes vor.
Genus CEDIOCERA Pascoe
Cediocera Pascoe, Ann. & Mag. Nat. Hist. (5) 20 (1887) 20.
CEDIOCERA TRISTIS Senna.
Cediocera tristis Senna, Not. Leyd. Mus. 14 (1892) 181.
Negros, Cuernos Mountains (Baker). Basilan (Baker).
Die Gattung umfasst 2 Arten, die aber nicht sicher trennbar
und die vielleicht nur Rassen einer Art sind. Es ist ubrigens
die einzige Gattung die einen mehr orientalischen Charakter hat.
Tristis hat eine grosse Migration und kommt bis Neu-Guinea
vor. Die urspriingliche Herkunft ist also noch immer erkenn-
bar. Mir lagen Belegstucke vor von: Malakka, Sumatra, Bor-
neo, Java, Neu-Guinea.
426 The Philippine Journal of Science ini
Genus HETEROPLITES Lacordaire
Heteroplites Lacordaire, Gen. Col. 7 (1866) 471.
HETEROPLITES ERYTHRODERES Boheman.
Heteroplites erythroderes Boheman, Schoenh. Gen. Cure. 5 (1840)
564.
Luzon, Provinz Ilocos Norte, Bangui (Banks) : Provinz Nueva
Vizcaya, Imugan (Boettcher) : Provinz Laguna, Mount Banahao
(Boettcher) : Provinz Camarines Sur, Mount Isarog, Balagbag
(Boettcher). Mindanao, Provinz Surigao, Surigao (Boett-
cher) : Provinz Bukidnon, Tangkulan (Boettcher) . Ballalon (?)
(Boettcher). Endemische Art. Farbung des Neu-Guinea
Typs. Eine zweite Art lebt auf Celebes mit Ausstrahlung nach
Borneo.
Genus DIURUS Pascoe
Diurus Pascoe, Journ. Ent. 1 (1862) 392.
DIURUS FURCILLATUS Gyllenhal.
Diurus furcillatus Gyllenhal, Schoenh. Gen. Cure. 1 (1833) 359.
Mindanao, Provinz Surigao, Surigao (Baker) : Provinz Da-
vao, Davao (Weber). Mindoro, Baco River (McGregor) .
Luzon, Ilocos Norte, Bangui (Banks). Samar (Baker).
Die gemeinste Art mit grosster Migration, Malakka, Sumatra,
Borneo, Java. Ueberall gleich haufig.
DIURUS PHILIPPINICUS Senna.
Diurus philippinicm Senna, Boll. Soe. Ent. Ital. 41 (1909) 45.
Endemische Art, naherer Fundort nicht angegeben.
DIURUS SAMARENSIS Kleine.
Diurus samarensis Kleine, Stett. Ent. Zeit. 87 (1926) 370, fig. 16.
Samar (Baker). Endemische Art.
DIURUS SHELFORDI Senna.
Diurus shelf ordi Senna, Proc. Zool. Soc. Lond. (1902) 279, t. 20,
fig. 6 ?.
Mindanao, Provinz Lanao, Kolambugan (Baker). Basilan
(Baker). Ausserdem von Borneo bekannt.
Diurus umfasst 25 Arten die mit geringen Ausnahmen orien-
talisch sind. In Neu-Guinea ist eine Art sicher nachgewiesen.
Auch auf den Carolinen soil eine vorkommen, es hat sie ausser
dem Autor wohl niemand gesehen. Diese Art bleibt unklar.
Auffallig ist das ganzliche Fehlen auf den Molukken, so dass
vorlaufig noch kein Anschluss der Orientalen an die Neu-Guinea-
46,
Kleine: Die Brenthiden der Philippinen-Inseln 427
Art moglich war. Vielleicht ist die Zuwanderung tiber die
kleinen Sunda-Inseln erfolgt und die Molukken sind tatsachlich
unberiihrt geblieben.
Was bei den Belopherini uber Herkunft und gegenwartige
Verteilung der Arten gesagt ist, tifft auch hier zu. Ja es ist die
Beteiligung des Ausgangszentrums. Neu-Guinea Molukken mit
den ostlichen Inselsehwarmen und Australien noch deutlicher als
bei den Belopherini. Von den 112 Arten sind 51 austro-malay-
isch oder australisch, 30 gehoren der orientalischen, 22 der neo-
tropischen Region an. Zu den Austromalayen diirfte eine Art
mit Ausstrahlung in die orientalische Region zu rechnen sein,
von einer ist der Fundort nicht sieher.
Mehrfach finden sich unter den wenigen Arten solche mit
Neu-Guinea-Farbung, also ganz analog den Verhaltnissen bei den
Belopherini.
Tabelle 8. — Verbreitungstabelle der Ithystenini.
Ma-
Iay-Hal-
binsel.
Suma-
tra.
+
+
Borneo.
Java.
Neu-
Guinea. ]
Achrionota bilineata Pascoe
+
+
+
+
+
+
i i i i i + i i
Achrionota spinifer Kleine a
Cediocera tristis Senna__
Heteroplites erythroderes Boheman a„ ._
Diurus furcillatus Gyllenhal
Diurm philippinicus Senna a
Diurus 8amaren8is Kleine a _ __
Diurus shelf or di Senna
a Endemisch.
PSEUDOCEOCEPHALINI
Genus OPISTHENOPLUS Kleine
Opisthenoplus Kleine, Deut. Ent. Zeit. 1 (1922) 139.
OPISTHENOPLUS CALABRESII Kleine.
Opisthenoplus calabresii Kleine, Arch. Nat. A. 10, 87 (1921) 35.
Luzon, Subprovinz Kalinga, Balbalasan (Boettcher) , mehr-
fach ohne naheren Fundort gesehen. Endemische Art mit
Ausfarbung des Neu-Guinea Typs.
OPISTHENOPLUS CAVUS F. Walker.
Opisthenoplus cavus F. Walker, Ann. & Mag. Nat. Hist. (3) 3 (1859)
262.
Luzon, Provinz Laguna, Mount Banahao (Boettcher) : Provinz
Nueva Vizeaya, Imugan (Boettcher). Mindanao, Provinz Su-
428
The Philippine Journal of Science
1931
rigao, Surigao {Boettcher) ; Provinz Lanao, Mumungan {Boett-
cher). Negros, Cuernos Mountains {Baker).
Haufige, weitverbreitete Art die in der ganzen orientalischen
Region vorkommt. Ich sah Belegstiicke von: Ceylon, Indien,
Burmah, Andamanen, Indo-China, Malakka, Sumatra, Borneo.
Hauptgebiet ist Indien. Von Java sah ich die Art noch nicht.
OPISTHENOPLUS FASCINATUS Kleine.
Opisthenoplus fascinatus Kleine, Deut. Ent. Zeit. 1 (1922) 140, figs.
7,8.
Luzon, Provinz Laguna, Mount Banahao {Baker) .
Die Verbreitung der Art ist wohl nur erst zum Teil bekannt.
Mir lagen belegstiicke vor von : Sumatra, Indien, Formosa. Das
Verbreitungszentrum lasst sich zur Zeit nicht angeben.
OPISTHENOPLUS PECUNDUS Kleine.
Opisthenoplus fecundus Kleine, Ent. Blatt. 19 (1923) 165.
Negros, Cuernos Mountains {Baker) . Endemische Art.
OPISTHENOPLUS MADENS Lacordaire.
Opisthenoplus modem Lacordaire, Gen. Col. 7 (1866) 455, nota 2.
Sibuyan (Sammler unbekannt).
Die Verbreitung von ma&ens hat einige Aehnlichkeit mit der
von cavus. Das eigentliche Zentrum muss aber mehr ostlieh lieg-
en, da mir noch niemals ein Belegstuck aus Indien vorgelegen
hat. Ich sah die Art von: Malakka, Andamanen, Indo-China,
Sumatra und Java. Die Ausbreitung nach Siidosten ist also auch
grosser als bei cavus.
Die 9 bekannten Arten sind orientalisch.
Genus HORMOCERUS Schoenherr
Hormocerus Schoenherr, Cure. Disp. (1826)
70.
HORMOCERUS RETICULATUS Fabricius.
Hormocerus reticulatus Fabricius, Syst. El.
2 (1801) 552.
Diese gemeine, von Ceylon bis Ost-Aus-
tralien iiberall vorkommende Art hat zahl-
reiche Rassen gebildet, die meist ineinan-
der iibergehen. Die von Boheman als
scrobicollis beschriebene Art kommt auf
den Philippinen vor und man findet in alteren Sammlungen hau-
fig zahlreiche Belegstiicke. Baker und Boettcher haben die Art
nicht gef unden. Es ist doch merkwiirdig, dass diese ausgezeich-
neten Sammler das Tier nicht gef unden haben.
Fig. 16. Verbreitungskarte
der Gattung Hormocerus
Schoenh.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 429
Genus APTERORRHINUS Senna
Apterorrhinus Senna, Not. Leyd. Mus. 17 (1895) 59.
APTERORRHINUS COMPRESSITARSIS Senna.
Apterorrhinus compressitarsis Senna, Not. Leyd. Mus. 17 (1895)
61.
Luzon, Provinz Ilocos Norte, Bangui (Boettcher). Siargao,
Dapa (Boettcher). Leyte (Boettcher).
Sehr weitverbreitete, aber nicht haufige Art. Ich sah Tiere
von: Malakka, Sumatra, Java, Buru.
APTERORRHINUS ALBATUS Kleine.
Apterorrhinus albatus Kleine, Arch. Nat. A. 3, 87 (1921) 226.
Luzon, Provinz Laguna, Mount Maquiling (Baker) .
Auch hier ist die Verbreitung noch ganz uniibersichtlich. Mir
lagen Belegstiicke von Neu-Guinea und Queensland vor.
Genus SCHIZOTRACHELUS Lacordaire
Schizotrachelus Lacordaire, Gen. Col. 7 (1866) 454.
SCHIZOTRACHELUS ANGULATICEPS Senna.
Schizotrachelus angulaticeps Senna, Boll. Soc. Ent. Ital. 31 (1899)
308.
Luzon, Subprovinz Kalinga, Balbalasan (Boettcher) : Provinz
Nueva Vizcaya, Imugan (Boettcher) : Provinz Ilocos Norte, Ban-
gui (Boettcher) . Negros, Cuernos Mountains (Baker) . Leyte,
Burauen (Boettcher). San Miguel (Boettcher). Basilan
(Baker) .
Weitere Funde sind bekannt von : Malayische Halbinsel, Bor-
neo, Celebes. Die Funde sind noch zu sporadisch und geben
keinen Ueberblick uber die Verbreitung.
SCHIZOTRACHELUS BAKERI Kleine.
Schizotrachelus bakeri Kleine, Arch. Nat. A. 10, 87 (1921) 33.
(a) Nominatform.
Luzon, Provinz Laguna, Los Baiios (Sammler unbekannt) :
Mount Banahao (Boettcher) : Subprovinz Benguet, Baguio
(Baker). Catanduanes, Virac (Boettcher). Negros, Orien-
tal Negros, Cuernos Mountains (Baker) : Occidental Negros, Fa-
brica (Schultze). Sibuyan (Baker). Polillo (Baker).
(b) Forma concolor.
Luzon, Provinz Laguna, Mount Banahao, Los Baiios (Baker) :
Provinz Nueva Vizcaya, Imugan (Boettcher). Mindanao, Su-
rigao, Surigao (Baker) : Provinz Lanao, Kolambugan, Iligan
(Baker). Siargao, Cabuntog (Boettcher). Negros (Baker).
430 The Philippine Journal of Science mi
Samar (Baker) . Basilan (Boettcher) . Endemische, aber wie
es seheint, auf alien Inseln vorkommende Art.
SCHIZOTRACHELUS BREVICAUDATUS Lacordaire.
Schizotrachelus brevicaudatus Lacordaire, Gen. Col. 7 (1866) 455,
nota 2.
Luzon, Provinz Laguna, Los Bafios (Sammler unbekannt).
Negros, Cuernos Mountains (Baker) . Sibuyan (Baker) .
Auf den grossen Sunda-Inseln ebenso haufig wie auf den Phil-
ippines
SCHIZOTRACHELUS BRUNNEUS Kleine.
Schizotrachelus brunneus Kleine, Arch. Nat. A. 10, 87 (1921) 36.
Luzon, Provinz Laguna, Los Baiios (Sammler unbekannt) :
Provinz Tayabas, Malinao (Baker) .
SCHIZOTRACHELUS CONSIMILIS Kleine.
Schizotrachelus conshnilis Kleine, Ent. Blatt. 19 (1923) 166.
Luzon, Provinz Laguna, Los Banos; Manila (Sammler unbe-
kannt). Ich sah die Art auch noch von Amboina.
SCHIZOTRACHELUS CORPULENTUS Kleine.
Schizotrachelus corpulentus Kleine, Arch. Nat. A. 10, 87 (1921)
32.
Mindanao, Provinz Agusan, Butuan (Baker). Endemische
Art.
SCHIZOTRACHELUS IMBRICELLUS Kleine.
Schizotrachelus imbricellus Kleine, Philip. Journ. Sci. 28 (1925) 604.
Luzon, Provinz Laguna, Mount Banahao (Baker). Endemi-
sche Art.
SCHIZOTRACHELUS IMITATOR Kleine.
Schizotrachelus imitator Kleine, Philip. Journ. Sci. 28 (1925) 603,
t. 1, fig. 10, 11.
Luzon, Subprovinz Kalinga, Balbalasan (Boettcher). Leyte,
Burauen, San Miguel (Boettcher). Panaon (Boettcher). Ca-
miguin (Boettcher). Catanduanes, Virac (Boettcher). En-
demische Art.
SCHIZOTRACHELUS INCONSTANS Kleine.
Schizotrachelus inconstans Kleine, Arch. Nat. A. 10, 87 (1921) 31.
Luzon, Provinz Laguna, Mount Maquiling (Baker).
Von den 31 Arten sind nur 8 nicht in der orientalischen Re-
gion. Von den 8 Nicht-Orientalen sind 6 auf den Molukken,
(einschliesslich Celebes) und nur 2 erreichen Neu-Guinea oder
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 431
Australien. Der Habitus der Gattung ist einheitlich, nur in der
Figur des Kopfes lassen sich zwei Gruppen erkennen. Diese
sind aber nicht zoogeographisch geschieden.
Von den 121 zur Tribus gehorenden Arten sind 33 der aethiop-
ischen, 23 der madegassischen und 43 der orientalischen Region
zuzuzahlen, 22 sind austro-malayisch bezeihungsweise austra-
lisch und 1 kommt von Ceylon bis Ost-Australien vor. Die Pseu-
doceocephalini gehoren also nicht dem Verbreitungskreis der
Belopherini und Ithystenini. Sie haben ihren Ursprung in Afri-
ka. Die Tribus hat einen intermediaren Charakter, die, wahr-
scheinlich aus den Trachelizini oder mit diesen aus einer noch
tieferen Wurzel entstanden, auch als Ausgangspunkt der Nemo-
cephalini anzusehen ist. Aus diesen haben sich die Brenthini
entwickelt, vielleicht sind auch die Taphroderini und Ulocerini
naher verwandt als man annimmt. Jedenfalls ist der Einfluss
auf die Brenthiden der neotropischen Region sehr gross. Nach
Osten hin ist der Einfluss geringer gewesen. Man vergleiche
hierzu meine Arbeit : "Die geographische Verbreitung der Bren-
thida>." a
Gesamt 267 Gattungen mit 1260 Arten. Davon sind im
Gebiet: Gesamt 48 Gattungen mit 124 Arten gefunden. Das
sind rund 18 Prozent der Gattungen und 9.9 Prozent der Arten.
Fur ein so kleines Verbreitungsgebiet eine stattliche Anzahl.
Von den Gattungen haben nur Miolispa und Schizotrachelus einen
grosseren Artbestand aufzuweisen. Sieben Tribus kommen
iiberhaupt nicht im Gebiet vor, sie sind mit Ausnahme von Eu-
trachelini auch nicht in der orientalischen, austromalayischen
und australischen Region vertreten.
Bestimmungstabelle der philippinischen Brenthiden.
A. TRIBUS
1. Russel in beiden Geschlechtern von gleicher Gestalt 2.
Prorostrum des c? von verschiedener Gestalt, aber niemals fadenformig,
des 9 immer lang, mehrfach so lang wie das Metarostrum, fadenfor-
mig, zum Bohren eingerichtet 4.
2. Prothorax am Halse verengt, zum Einlegen der Voderbeine eingerich-
tet, Elytren am Hinterrand an der Sutura zugespitzt... Calodromini.
Prothorax nicht verengt, Elytren nicht zugepitzt 3.
3. Tibien der Vorderbeine mit grossem Innenzahn Stereodermini.
Tibien ohne Innenzahn Trachelizini.
4. Kopf und Russel, oder wenigstens der letztere, deformiert.
Amorphocephalini.
Nicht deformiert 5.
aiArch. Nat. A. 10, 87 (1921) 38-132.
432
The Philippine Journal of Science
Tabblle 9. — Verbreitungstabelle der Pseudoceocephalini.
1931
Opisthenoplus calabresii Kleine*.
Opisthenoplus cavus F. Walker. _
Opisthenoplus fascinatus Kleine__
Opisthenoplus fecundus Klein e »._
Opisthenoplus madens Lacor-
daire
Hormocerus reMculatus Fabri-
cius
Apterorrhinus compressitarsis
Senna
Apterorrhinus albatus Kleine
Schizotrachelus angukkticeps
Senna
Schizotrachelus bakeri Kleine »___
Schizotrachelus brevicaudatus
Lacordaire
Schizotrachelus brunneus Kleine a
Schizotrachelus consimilis Kleine
Schizotrachelus corpulentus
Kleine a
Schizotrachelus imbricellus
Kleine a
Schizotrachelus imitator Kleine a.
Schizotrachelus inconstans
Kleine a
a Endemisch.
(b)
(b)
(b)
3*
+
cb)
(b)
cb)
(*>
b Von Australien bis Ceylon gemein.
5. Fiihler vor der Mitte des Riissels eingefugt, Metatarsus auffallend ver-
langert Ithystenini.
Fiihler in der Mitte des Riissels stehend, Metatarsus nicht auffallend
verlangert 6.
6. Fiihler und Beine lang, Habitus schlank Belopherini.
Fiihler und Beine kurz, Habitus gedrungen 7.
7. Mandibeln das Mannes gross, Riissel gedrungen Arrhenodini.
Mandibeln das Mannes klein, Riissel schlank, mehr oder weniger walzig.
Pseudoceocephalini.
B. GATTUNGEN
1. CALODROMINI
1. Tarsen hypermorph, Metatarsus fast so lang wie das ganze Tier.
Calodromus Guerin.
Tarsen im Verhaltnis zum Korper und den Beinen normal, hochstens
der Metatarsus so lang wie das 2. und 3. Glied zusammen 2.
46,3
Kleine: Die Brenthiden der Philippinen-Inseln 433
Tabelle 10. — Starkenverhaltnis der philippinischen Gattungen und Arten
zu den Brenthiden der ganzen Welt.
Calodromini
Stereodermini
Trachelizini
AmorphocephalinL _ .
Arrhenodini
Belopherini
Eutrachelini
Tychaeini
Ithystenini
Ulocerini
PseudoceocephalinL
Taphroderini
Rhyticephalini
Nemocephalini
Brenthini
Der ganzen | Davon im
Welt. ! Gebiet.
gen.
Arten.
62
181
8
89
27
225
17
83
41
196
24
95
1
1
1
1
21
108
2
22
39
124
8
32
! *
2
12
3
5.
6.
2. Hinterschienen hypermorph oder doch von auffallender Gestalt, niemals
normal 3.
Hinterschienen normal 4.
3. Auf den Elytren sind alle Rippen gleichmassig entwickelt.
Cyphagogus Parry.
Zweite Rippe verjsiirzt Epigogus Kleine.
4. Fuhler nach vorn in grossen Gruben stehend, die durch einer mehr
oder weniger schmale Wand getrennt sind 5.
Fuhler seitlich stehend, in Riisselbreite getrennt 6.
Auf den Elytren ist die 2. Rippe weit unterbrochen.
Orthopareia Kleine.
Die 3., 5. und 7. Rippen vekurzt Asaphepterwn* Kleine.
Prothorax vorn gar nicht oder nur ganz wenig verengt 7.
Prothorax vorn immer zum Einlegen der Beine verengt 9.
7. Unterseite des Kopfes oder Riissels ohne Zahn oder buckliger Ver-
diekung auf den Seitenkanten Opisthenoxys Kleine.
Unterseite mit mehr oder weniger grossem Zahn oder buckliger Ver-
dickung auf Kopf oder Metarostrum 8.
8. Korper schuppenartig breit behaart Pseudo cyphagogus Desbr.
Korper nicht schuppenartig behaart Mesoderes Senna.
9. Kopf unterseits nicht gezahnt Atopomorphus Kleine.
Kopf unterseits gezahnt 10.
10. Auf den Elytren sind die 1. und 3. Rippen an der Basis verkiirzt, von
der 2. und 4. eingeschlossen Dictyotopterus Kleine.
Alle Rippen normal lang Eterozemup Senna.
263774 9
434 The Philippine Journal of Science ij>sj
2. STEREODERMINI
1. Fuhler sehr lang und diinn, zuweilen von Korperlange.
Jonthocerus Lacordaire.
Fuhler kurz, gedrungen 2.
2. Neunte bis elfte Fiihlerglied verdickt, erheblich grosser als die vor-
hergehenden Stereo dermus Lacordaire.
Neunte bis elfte Fiihlerglied nicht verdickt, zuweilen kaum so lang wie
die vorhergehenden Cerobates Schoenherr.
3. TRACHELIZINI
1. Elytren mit erhabenen und tiefliegenden Rippen, Hinterrand an der
Sutura verlangert Hoplopisthius Senna.
Elytren mit gleichhohen Rippen, Hinterrand gerundet 2.
2. Prothorax ungefurcht 3.
Prothorax gefurcht 4.
3. Auf den Elytren sind alle Rippen ausgebildet.... Miolispa Pascoe (pars).
Nur die Sutura ist voll entwickelt, die folgenden Rippen fehlen ganz
oder sind rudimentar Homophylus Kleine.
4. Vorderschienen innenseits keilformig erweitert Metatrachelizus Kleine.
Vorderschienen nicht erweitert 5.
5. Ausser der Sutura sind nur noch erne bis zwei Rippen vorhanden.
Trachelizus Schoenherr.
Alle Rippen sind entwickelt 6.
6. Kopf oberseits und an den Seiten mehrfach eingekerbt, oder tuberkel-
artig verdickt 7.
Kopf nicht eingekerbt, glatt gerundet Microtrachelizus Senna.
7. Kopf langer als breit, Augen nach vorn geruekt Miolispa Pascoe.
Kopf kurz, Augen immer an der Basis stehend 8.
8. Grossere Arten, 10 bis 15 mm., Schenkel wehrlos- Hypomiolispa Kleine.
Kleine Arten, 5 bis 8 mm., vor der Schenkel gedornt Higonius Lewis,
4. AMORPHOCEPHALINI
1. Kopf rundlich, nur der Riissel deformiert Cordus Schoenherr.
Kopf und Riissel deformiert.. 2.
2. Prorostrum schmaler als das Metarostrum, unterseits glatt, nicht ge-
rinnt Leptamorphocephalus Kleine.
Prorostrum breiter als das Metarostrum, unterseits mehr oder weniger
vorgezogen oder verdickt Par amor phocephalus Kleine.
3. ARRHENOD1NI
1. Riissel so breit wie der Kopf oder kaum schmaler, robust, Mandibeln
immer kraftig, einen freien Raum einschliessend oder nicht 2.
Riissel schmaler als der Kopf, nach dem Vorderrand erweitert, mit
grossen, vorstehenden Mandibeln die einen grossen, freien Raum
einschliessen Eupsalis Lacordaire.
Riissel von ahnlicher Gestalt, Mandibeln klein 5.
Riissel am Vorderrand nicht erweitert, mehr oder weniger parallel,
Mandibeln klein bis sehr klein Amphicordus Heller.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 435
2. Prorostrum sehr breit, mehr oder weniger parallel, Vorderrand tief ein-
geschnitten und die Mandibeln im Einschnitt verborgen.
Agriorrhynchus Power.
Prorostrum gegen den Vorderrand nur gerung verbreitert, Mandibeln
nicht in einer Einbuchtung des Vorderrandes verborgen, sondern vor-
stehend 3.
3. Riissel dick, walzig, im Verhaltnis zum Kopf sehr lang.
Eupeithes Senna.
Riissel nicht walzig, normal lang 4.
4. Mandibeln gross, zangenartig, einen freien Raum einschliessend, Kopf
meist sehr lang, Augen klein, nach vorn geriickt.
Prophthalmus Lacordaire.
Mandibeln nicht gross, nicht zangenartig, nur einen kleinen, freien
Raum einschliessend Baryrrhynchus Lacordaire.
5. Kopf hinter den Augen seitlich gedornt oder uber den Hals nach hinten
vorgezogen Csenorychodes Kleine.
Kopf nicht gedornt oder uber den Hals erweitert Pseadowjchodes Senna.
6. BELOPHERINI
1. Prorostrum am Vorderrand gar nicht oder nur gering verbreitert, je-
denfalls nicht nach den Seiten ausladend 2.
Prorostrum am Vorderrand nach den Seiten spitz verbreitert 3.
2. Schenkel ungedornt Ypselogonm Kleine.
Schenkel gedornt Heteroblysmia Kleine.
3. Metarostrum mit starkem Seitenzahn Apocemns Calabresi.
Ohne Seitenzahn 4.
4. Mandibeln sehr gross, einen freien Raum einschliessend.
Henarrhenodes Heller.
Mandibeln klein, keinen freien Raum einschliessend.
Anepsiotes Kleine.
7. ITHYSTENINI
1. Elytren glatt, ausser der Sutura hochstens noch eine Rippe vorhan-
den, die folgenden nur punktstreifig 2.
Elytren regelmassig punktstreifig, neben der Sutura keine scharfen
Rippen 3*
2. Erste und zweite Abdominalsegment deutlich gefurcht.
Cediocera Pascoe.
Abdomen nicht gefurcht, hochstens schwach abgeplattet.
AchrionotaJ Pascoe.
3. Ohne kleiige Beschuppung, Prothorax gefurcht Heteroplites Lacordaire.
Mit kleiiger Beschuppung, Prothorax ungefurcht Dntrus Pascoe.
8. PSEUDOCEOCEPHALINI
1. Elytren am Hinterrand mehr oder weniger verlangert.
Opisthenophis Kleine.
Elytren nicht verlangert 2-
2. Elytren an der Basis ungezahnt Schizo tracheitis Lacordaire.
Elytren an der Basis gezahnt - 3.
3. Klauenglied wenn auch kraftig, so doch keulig Hormocerus Schoenherr.
Klauenglied walzig, seitlich zusammengepresst... Apterorrhiwus Senna.
436 The Philippine Journal of Science 1931
c. AKTEN
1. CALODROMINI
Genus CALODROMUS Guerin
Metatarsus mit einem Zahn C. mellyi Guerin.
Mit zwei Zahnen C. crinitus Kleine.
Genus CYPHAGOGUS Parry
1. Elytren mit 2 rotgelben Binden auf jeder Seite.
C modiglianii Senna.
Elytren einfarbig, schwarz 2.
2. Pro- und Mesorostrum und eine kielformige Platte auf dem Metaros-
trum glanzend; Kopf und Riissel sonst matt . 3.
Kopf und Riissel gleichmassig glatt 4.
3. Der glanzende Teil des Riissels zart punktiert; 1. und 2. Abdominal-
segment nicht gefurcht C. planifrons Kirsch.
Der glanzende Teil an der Basis grob, rugos punktiert; 1. und 2. >Ab-
dominalsegment keilformig, kraftig gefurcht.... C. gladiator Kleine.
4. Kopf iiber den Augen mit groben, zuweilen zu einer Furche verschmolz-
enen Punkten; Kopf grob, einzeln punktiert, in den Punkten be-
haart 5.
Kopf ohne Augenfurche, unbehaart, selten mit einzelnen Harchen am
Hinterkopf 6.
5. Unterseite des Kopfes mit mehreren Querwiilsten.
C. longulus Senna.
Ohne Querwiilste — C. silvanus Senna.
6. Metatarsus der Hinterbeine ohne Stiel kiirzer als die 2. und 3. Glieder
zusammen 7.
Metatarsus langer als die 2. und 3. zusammen 8.
7. Schlanke Art, Thoracalconus bucklig, Stiel der Hinterschenkel an der
Keule unterseits tief, fast halbkreisformig eingekerbt.
C. westwoodi Parry.
Gedrungene Art, Thoracalconus rechwinklig, gerade aufsteigend, Stiel
der Hinterschenkel an der Keule nicht eingekerbt.
C. buccatus Kleine*
8. Stiel der Hinterschenkel gerade, am Uebergang zur Keule nicht ve-
rengt oder auf Ober- und Unterseite eingekerbt 9.
Stiel der Hinterschenkel an der Keule verengt oder eingekerbt 10.
9. Riissel schmal, viel langer als der Kopf C tabacicola Senna.
Riissel nicht auffalend verschmalerfc, so lang oder kiirzer als der Kopf.
C. simulator Senna.
10. Untere Halfte der Ftihler; Wurzel der Schenkel und die drei letzten
Abdominalsegmente rotlich C. eichhomi Kirsch.
Einfarbig schwarz C. whitei Westwood.
Genus EPIGOGUS Kleine
Nur eine Art E. flexibilis Kleine.
Genus ORTHOPAREIA Kleine
Nur eine Art O. idonea Kleine.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 437
Genus ASAPHEPTERUM Kleine
JNJur erne Art j± fovmosanum Kleine.
Genus OPISTHENOXYS Kleine
Fiihlerglied am langsten, alle Glieder langer als breit.
O ochvcLCGii^s xCleinp
Fiihlerglied am langsten, die folgenden Glieder perlig.
0. hoettcheri Kleine.
Genus PSEUDOCYPHAGOGUS Desbr.
Nur eine Art Pt squamifer Desbr.
Genus MESODERES Senna.
Nur eine Art Mt fessus Kleine>
Genus ATOPOMORPHUS Kleine
Nur eine Art A% schulUei Kleine.
Genus ETEROZEMUS Senna
Elytren rotbraun, Querbinde schwarz E. pubens Senna.
Elytren schwarz mit 4 braunen Makeln E. Isetus Senna.
Genus DICTYOTOPTERUS Kleine
Zweifarbige Art; Prothorax, Kopf und Riissel ziegelrot; Elytren blau-
schwarz; Prothorax unbehaart D. pulcherrimus Kleine.
Einfarbige Art; Prothorax, namentlich an den Seiten, zottig behaart.
D. philippinensis Kleine.
2. STEREODERMINI
Genus JONTHOCERUS Lacordaire
1. Zweite Rippe der Elytren in der Mitte nicht unterbrochen.
J. latico statis Kleine.
Zweite Rippe mehr oder weniger, meist betraehtlich unterbrochen.... 2.
2. Prothorax ungefurcht, rot gefarbt; Elytren von tiefschwarzer Farbe.
J. bicolot Heller.
Prothorax kraftig gefurcht oder an der Basis tief grubig eingedriickt,
das ganze Tier einfarbig 3
3. Kopf hinter den Augen bestimmt winklig J. modiglianii Senna.
Kopf hinter den Augen gerundet J. asiaticus Kleine.
Genus STEREODERMUS Lacordaire
Nur eine Art S. flavotibialis Kleine.
Genus CEROBATES Schoenherr
1. Aussenecken der Elytren am Absturz kurz gezahnt.
C. clinatus Kleine.
Aussenecken gerundet 2.
2. Prothorax ungefurcht 3.
Prothorax gefurcht 7.
438 The Philippine Journal of Science 1931
3. Elytren von der Sutura bis zum Absturz dreifurchig, an den Seiten
leicht gestreift oder schwach punktiert; 3. Furehe bis zum Absturz
verlangert, zuweilen in der Mitte obsolet seltener verschwommen und
unsicher 4.
Elytren nur an der Basis dreifurchig, an den Seiten glatt oder leicht
gestreift, 3. Furehe immer gegen den Absturz verschwindend 6.
4. Elytren gegen den Absturz lang, auffallig verschmalert.
C. angustipennis Senna,
Elytren im Apicalteil normal verschmalert 5.
5. Prorostrum bestimmt langer als das Metarostrum... C. sequalis Kleine.
Pro- und Metarostrum gleichlang C. tristriatus Fabricius.
6. Kleine Art; Riissel robust; Kopf hinter den Augen gerundet, Seiten
der Elytren glanzend C. sexsulcatus Motschulsky.
Grosse Art; Riissel zart; Kopf hinter den Augen mehr oder weniger
winklig; Seiten der Elytren gestreift C. adustus Senna.
7. Kopf oberseits iiber den Hals zuriickgezogen ; innen dreieckig eingekerbt.
C. costatus Kleine.
Oberseite des Kopfes gerade oder nur flaeh nach innen gebuchtet.... 8.
8. Elytren mit durchgehender dritter Furehe C. grouvellei Senna.
Elytren mit verkxirzter 3. Furehe 9.
9. Furehe des Prothorax durchgehend, 1. Fiihlerglied langlich.
C. sumatranus Senna.
Furehe nur in der basalen Halfte, 11. Fiihlerglied kurz.
C. formosanus Schonf.
TRACHELIZINI
Genus HOMOPHYLUS Kleine
Nur eine Art H. mindanensis Kleine.
Genus METATRACHELIZUS Kleine
Nur eine Art M. constans Kleine.
Genus TRACHELIZUS Schoenherr
Nur eine Art T. bisuldatus Fabricius.
Genus MIOLISPA Pascoe
1. Elytren nur auf dem Absturz gerippt-gefurcht, sonst glatt und nur
zart punktiert 2.
Elytren auf der ganzen Flache gerippt-gefurcht 3.
2. Einfarbig schwarze Art M. paucicostata, Kleine.
Kopf, Fiihler und Riissel schwarz; Prothorax zinnoberrot; Elytren
blaumetallisch M. pulchella Kleine.
3. Prothorax deutlich und kraftig langsgefurcht 4.
Prothorax ganz obsolet oder ungefurcht 6.
4. Prothorax iiberall dicht und tief grubig punktiert 5.
Prothorax hochstens am Hinterrande in sehr geringem Umfange oder
gar nicht punktiert M. fraudatrix Kleine.
5. Einfarbige matte, schwarze Art, 3. Rippe nicht gelb.
M. unicolor Kleine.
Grunlich-erzfarben, glanzend, 3. Rippe gelb M. persimilis Kleine.
6. Prothorax mindestens im basalen Teil deutlich und kraftig punktiert 7.
Prothorax unpunktiert, hochstens am Hinterrande mit engen Punkten 17.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 439
7. Prothorax rot; Elytren schwarz 8.
Farbe der Elytren und des Prothorax ubereinstimmend oder die Ely-
tren hell und der Prothorax dunkel gefarbt 9.
8. Matte Art, 3. Rippe der Elytren nicht gelb M. bicolor Kleine.
Hochglanzende Art, 3. Rippe gelb M. clavicornis Kleine.
9. Schenkel der Mittel- und Hinterbeine verdickt, gross, klobig, Stiel kurz,
breit, zusammengedriickt 10.
Schenkel normal, keulig, Stiel diinn, deutlich abgesetzt 11.
10. Kopf und Riissel mit Ausnahme des vorderen Prorostrums matt, Kor-
perseiten dunkel gefarbt M. flexilis Kleine.
Am ganzen Korper hochglanzend, Korperseiten hellrotbraun.
M. fornicata Kleine.
11. Schienen aller Beine, namentlich der Vorder- und Hinterbeine mit
starkem Innenzahn M. formosa Kleine.
Schienen normal ohne Zahn 12.
12. Rotbraune Arten, 3. Rippe gelb oder nicht 13.
Dunkelfarbige Arten, 3. Rippe immer gelb M. flavolineata Kleine.
13. Prothorax matt 14.
Prothorax hochglanzend 15.
14. Furche des Metarostrums zu einer sammetartigen, matten Platte verei-
nigt M. lineata Senna.
Nicht sammetartig, nicht vereinigt M. siporana Senna.
15. Fuhlerglieder eng stehend, 2. Glied breiter als alle anderen.
M. robusta Kleine.
Fiihler normal 16.
16. Paramerenlamellen fingerartig, ausser der Sutura eine deutliche
schwarze Makel hinter der Elytrenmitte..... M. cruciata Senna.
Paramerenlamellen weit getrennt, zangenartig, nur die Sutura dunkel,
Makel meist fehlend, seltener unscharf vorhanden.
M. intermedia Senna.
17. Elytren ausser der Sutura auf der ganzen hinteren Halfte schwarz.
M. ephippium Kleine.
Elytren rotbraun, hochstens die Sutura and eine oder zwei Makeln ver-
dunkelt 18
18. Nur die Sutura ist dunkel gefarbt M. pascoei Kleine.
Ausser der Sutura ist noch eine dunkle Makel vorhanden 19.
19. Mit Ausnahme des 3. sind alle Fuhlerglieder breiter als lang.
M. discors Senna.
Mit Ausnahme des 2. sind alle Fuhlerglieder langer als breit.
M. elongata Kleine.
Genus HYPOMIOLISPA Kleine
1. Neunte und zehnte Fiihlerglied lang, walzig, zylindrisch, mehrfach so
lang wie die vorhergehenden, 11. so lang wie das 9. und 10. zusam-
men 2.
Neunte und zehnte Fiihlerglied zwar langer als die vorhergehenden,
aber niemals zylindrisch, sondern tonnenformig, kurz, mehr oder
weniger rundlich oder fast quadratisch, 11. meist kurz, zuweilen nur
wenig langer als das 8. oder 9 3.
2. Unterseite vom Prothorax bis zum Abdomen an den Seiten mit sil-
berglanzenden Flecken H. exarata Desbr.
440 The Philippine Journal of Science *93i
Ohne diese Flecken; Kopf, Riissel und Unterkanten der Schenkel mit
starkem Tomment bedeckt, sonst glatt H. tomentosa Kleine.
3. Schlanke, kleine Formen, Kopf mehr oder weniger quadratisch-eckig
oder etwas langer als breit; Prothorax mit seinen Organen immer
schwarz; Elytren niemals quer-schwarzstreifig 4.
Kopf mehr oder weniger robust, zuweilen gedrungen oder grossere
Formen mit querem dreieckigem, niemals viereckigem Kopfe; Pro-
thorax nur bei einigen Arten schwarz und dann ist die Grundfarbe
iiberhaupt schwarz; die Elytren sind rot gezeichnet und die Beine
sind bunt oder die Elytren sind querstreifig 5.
4. Der hinter den Augen liegende Teil des Kopfes ist doppelt so gross
wie der Augendurchmesser, Penis mit kurzen Parameren.
H. nupta Senna.
Der hinter den Augen liegende Teil hochstens so gross wie der Augen-
durchmesser; Parameren sehr lang H. sponsa Kleine.
5. Hinterrand des Kopfes an den Seiten nicht gezahnt, Augen gross, den
Hinterrand beriihrend H. ocularis Kleine.
Kopf hinter den Augen gezahnt, die Augen den Hinterrand nicht be-
riihrend 6.
6. Prothorax schwarz, Elytren mit 2 schwarzen Querbinden.
H. trachelizoides Senna.
Prothorax rotbraun, Elytren nicht mit schwarzen Banderungen, son-
dern mit verdunkelter Sutura und gleichem Aussenrand, zuweilen
mit undeutlichen Makel auf der Mitte der Sutura.
H. helleri Kleine.
Genus HIGONIUS Lewis
Nur eine Art H. cilo Lewis.
Genus MICROTRACHELIZUS Senna
1. Metarostrum 3-furchig 2.
Metarostrum 1-furchig 3.
2. Kopf oberseits ungefurcht M. pubescens Senna.
Kopf oberseits gefurcht M. tabaci Senna.
3. Auf den Elytren ist die 2. Rippe nur am Absturz vorhanden, 3. dureh-
gehend, verdickt, 4. und 5. verkiirzt, 6. am Absturz verdickt, 7. normal
durchgehend, die ubrigen fehlen ganz M. fluxus Kleine.
4. Rippe nicht unterbrochen, wenn auch verschmalert, keine Rippe ist
verkiirzt und keine fehlt M. siamensis Kleine,
Genus HOPLOPISTHIUS Senna
Nur eine Art H. trichimerus Senna.
AMORPHOCEPHALINI
Genus CORDUS Schoenherr
Nur eine Art C. peguanus Senna.
Genus LEPTAMORPHOCEPHALUS Kleine
Nur eine Art L. feeder atus Kleine.
Genus PARAMORPHOCEPHALUS Kleine
Nur eine Art P. setosus Kleine.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 441
ARRHENODINI
Genus AGRIORRHYNCHUS Power
Nur eine Art A. ignarius Kleine.
Genus EUPEITHES Senna
Nur eine Art E. dominator Kleine.
Genus PROPHTHALMUS Lacordaire
Prothorax mit tiefer Mittelfurche P. longirostris Gyllenhal.
Prothorax ungefurcht P. tricolor Power.
Genus BARYRRHYNCHUS Lacordaire
Nur eine Art B. schroederi Kleine.
Genus EUPSALIS Lacordaire
Nur eine Art E. kleinei Heller.
Genus CAENORYCHODES Kleine
Sehmuckstreifen auf den Elytren lang, 3. Rippe basal bis zur Mitte
und mit kurzer Unterbrechung oder durchgehend bis zum Absturz
verlangert, 4. wie die 3. oder ahnlich, niemals in kurzen Querbinden.
C. splendens Kirsch.
Sehmuckstreifen nicht lang, sondern in kurzen Querbinden angelegt.
C. serrirostris Fabricius.
Genus PSEUDORYCHODES Senna
Nur eine Art P. prmclarus Kleine.
Genus AMPHICORDUS K. M. Heller
Nur eine Art A. improportionalis Heller.
BELOPHER1NI
Genus YPSELOGONIA Kleine
Nur eine Art Y. peregrina Kleine.
Genus HETEROBLYSMIA Kleine
1. Prothorax glatt, ohne rugosem Fleck auf der Mitte.
H. formidolosa Kleine.
Mit rugosem Fleck 2.
2. Violettbraun, Prothorax ziegelrot, Schmuchzeichnung auf den Elytren
langsstreifig H. electa Kleine.
Einfarbig braun, Schmuckzeichnung nicht langstreifig.
H. accurata Kleine.
Genus APOCEMUS Calabresi
Nur eine Art A. ignobilis Kleine.
Genus HENARRHENODES K. M. Heller
Nur eine Art H. macgregori K. M. Heller.
Genus ECTOCEMUS Pascoe
Nur eine Art #. badeni Kirsch.
442 The Philippine Journal of Science 1931
Genus ANEPSIOTES Kleine
Kopf breiter als lang; Prothorax an den Seiten mit schwarzen Makeln.
A, luzonicus Calabresi.
Kopf quadratisch ; ohne schwarze Makeln A. nitidicollis Calabresi.
ITHYSTENINI
Genus CEDIOCERA Pascoe
Nur eine Art C. tristis Senna.
Genus ACHRIONOTA Pascoe
Am ganzen Korper in der Punktierung schuppig behaart.
A. bilineata Pascoe.
In den Punkten nicht behaart A. sjrinifer Kleine.
Genus HETEROPLITES Lacordaire
Nur eine Art H. erythroderes Boheman.
Genus DIURUS Pascoe
1. Basales Fiihlerglied hoehstens massig verlangert und an der Spitze
niemals nodos verdickt 2.
Basales Fiihlerglied immer lang, schlank, an der Spitze nodos verdickt.
3.
2. Elytren in beiden Geschlechtern nur gedornt, Spitzenglieder der Fiih-
ler getrennt D. shelfordi Senna.
Elytren mit deutlichen Anhangen, nicht gedornt, Spitzenglieder der
Fiihler dich stehend D. furcillatus Gyllenhal.
3. Zweite Fiihlerglied h so lang wie das 3 D. philippinicus Senna.
Zweite und dritte Fiihlerglied gleichlang D. samarensis Kleine.
PSEUDOCEOCEPHALINI
Genus OPISTHENOPLUS Kleine
1. Hinterer Augenrand deutlich gezahnt 2.
Augenrand ungezahnt oder flach gekerbt 4.
2. Augenrand mit 3 Zahnen O. fascinatus Kleine.
Augenrand mit 2 Zahnen 3.
3. Pechbraun; Tarsen der Hinterbeine des <$ walzig, Prothorax an der
Basis kraftig punktiert 0. madens Lacordaire.
Rotbraun, Tarsen der Hinterbeine des c? kegelig, Prothorax garnicht
oder wenig punktiert O. cavus F. Walker.
4. Schwarz, Prothorax ziegelrot O. calabresii Kleine.
Einfarbig hellrotbraun O. fecundus Kleine.
Genus HORMOCERUS Schoenherr
Nur eine Art H. reticulatus Fabricius.
Genus APTERORRHINUS Senna
Schwarz, nur die Tarsen rotbraun, Kopf nicht gefurcht, Suturalfurche
gegittert A. albatus Kleine.
Rotbraun, Kopf tief gefurcht, Suturalfurche nur punktiert, nicht ge-
gittert A. compressitarsw Senna.
46,3 Kleine: Die Brenthiden der Philippinen-Inseln 443
Genus SCHIZOTRACHELUS Lacordaire
1. Kopf etwa quadratisch oder wenig langer als breit 2.
Kopf schmal, viel langer als breit, parallel oder oblong 3.
2. Schwarze Art S. brevicaudatus Senna.
Kirschrote, zuweilen etwas dunklere Art S. brunneus Kleine.
3. Zweifarbige Art, Prothorax rot, sonst violettbraun (Nominatform).
S. bakeri Kleine.
Einfarbige Arten 4.
4. Pechschwarze Art, Elytren am Hinterrand in der Mitte eingeschnitten.
5.
Heller oder dunklerbraune Arten 7.
5. Prothorax tief punktiert, Elytren am Absturz mit stark verdickter 8.
Rippe S. imbricellus Kleine.
Prothorax unpunkfciert oder nur mit einigen zarten Punkten, Elytren
nicht mit verdickter 8. Rippe 6.
6. Kopf gegen den Russel verengt, Meta- und Prorostrum schmal gefurcht,
Hinterschienen breit S. angulaticeps Senna.
Kopf parallel; Metarostrum breit und tief gefurcht; Prorostrum ohne
Purche; Hinterschienen schmal 5. imitator Kleine.
7. Prothorax tief punktiert S. inconstans Kleine.
Prothorax unpunktiert 8.
8. Hinterrand des Kopfes tief dreieckig eingekerbt.
S. bakeri Kleine f . concolor.
Hinterrand des Kopfes breit, flach eingekerbt 8.
9. Schwarzbraun, Kopf zart punktiert S. corpulentus Kleine.
Rotbraun, Kopf unpunktiert S. consimilis Kleine.
ILLUSTRATIONEN
KARTEN
Fig. 1. Verbreitungskarte der Gattung Calodromus Guer.
2. Verbreitungskarte der Gattung Cyphagogiis Parry.
3. Verbreitungskarte der Gattung Oplsthenoxys Kleine.
4. Verbreitungskarte der Gattung Mesodei*es Senna.
5. Verbreitungskarte der Gattung Jonthocerus Lacord.
6. Verbreitungskarte der Gattung Stereodermus Lacord.
7. Verbreitungskarte der Gattung Cerobates Schoenherr.
8. Verbreitungskarte der Gattung Metatrachelizus Kleine.
9. Verbreitungskarte der Gattung Miolispa Pascoe.
10. Verbreitungskarte der Gattung Microtrachelizus Senna.
11. Verbreitungskarte der Gattung Cordus Schoenh.
12. Verbreitungskarte der Gattung Baryrrhynchus Lacord.
13. Verbreitungskarte der Gattung Eupsalis Lacord.
14. Verbreitungskarte der Gattung Caenorychodes Kleine.
15. Verbreitungskarte der Gattung Pseudorychodes Senna.
16. Verbreitungskarte der Gattung Hormocerus Schoenh.
445
NEW OR LITTLE-KNOWN TIPULID^E FROM THE
PHILIPPINES (DIPTERA), XIP
By Charles P. Alexander
Of Amherst, Massachusetts
TWO PLATES
The crane flies discussed in the present report are all from
Davao district, Mindanao, Philippine Islands, where they were
collected by my friend and former student Mr. Charles F. Clagg.
The majority of the specimens were taken at high altitudes on
Mount Apo, which was twice ascended to the summit by Mr.
Clagg. Other species from this rich collection will be discussed
in later parts under this general title. All types are preserved
in the author's collection.
LIMONIIN^E
LIMONIINI
LIMONIA (LAOSA) MANOBO sp. nov. Plate 1, fig. 1.
Ground color of notum whitish, the prsescutum with four
chestnut-brown stripes; femora yellow, the tips broadly black-
ened ; wings whitish, with an irregularly banded yellow pattern,
the areas bordered by darker; the supernumerary crossvein in
cell R3 lying far distad of the one in cell R5.
Male. — Length, about 7.5 millimeters; wing, 9.5.
Rostrum and palpi black, the former about one-half the re-
mainder of the head. Antennae with the scapal segments black;
first flagellar segment light yellow, the remaining segments pass-
ing through brown to black ; flagellar segments oval, clearly de-
marked, each with one seta that is a little longer than the seg-
ment, unilaterally arranged, in addition to several small setse;
terminal segment one-half longer than the penultimate, the ter-
minal two setae small. Head brownish gray, the center of the
posterior vertex narrowly blackened, the narrow anterior vertex
light golden yellow.
1 Contribution from the entomological laboratory, Massachusetts State
College.
447
448 The Philippine Journal of Science 1931
Pronotum medially obscure yellow, dark brown sublaterally.
Mesonotal prsescutum with the restricted ground color whitish,
the disk almost covered by four confluent chestnut-brown stripes
that are narrowly bordered by blackish, the lateral stripes con-
tinued laterad to the margin, leaving a large humeral area of the
ground color completely isolated from a small area before the
suture ; scutal lobes light orange, bordered by blackish, the median
area darkened ; scutellum yellow, the caudal portion with a large
brown spot ; postnotal mediotergite chiefly dark brown, the cepha-
lic portion more yellowish, especially medially. Pleura whitish,
extensively variegated with dark brown, the major areas includ-
ing most of the anepisternum and sternopleurite, together with
the pleurotergite, and a small spot on the pteropleurite. Halteres
black, the base and apex of the stem narrowly and subequally
light yellow. Legs with the coxae pale yellowish white, varie-
gated with brown; trochanters yellow; femora yellow, the tips
very broadly blackened, the amount including about the distal
quarter and subequal in amount on all legs ; tibise light yellow ;
tarsi yellow, the outer segments blackened. Wings (Plate 1,
fig. 1) whitish, with an irregularly banded brownish yellow
pattern that is suggestive of that of many species of Epiphragma;
the bands include a restricted postarcular area ; a complete band
at near midlength of cells R and M, widened out along vein Cu,
ending at margin at vein 2d A ; bands at cord and outer end of cell
1st M2, broadly confluent in the stigmal region, the latter ex-
tended out across the supernumerary crossveins in the radial
field to the margin at midlength and apex of cell R2 ; all bands
margined with brown; an isolated small brown spot at end of
vein 1st A; cells C and Sc uniformly darkened. Venation: Sct
ending beyond r-m, Sc2 close to its tip ; R, bent strongly caudad
at R2 ; supernumerary crossvein in cell R3 lying far more than its
own length beyond the one in cell R5; second section of M1 + 2
strongly sinuous ; m-cu about one-half its length beyond the fork
of M; cell 2d A wide.
Abdominal tergites dark brown, narrowly pale medially and
sublaterally at base; hypopygium chiefly darkened. Male hypo-
pygium almost as in the typical form of the subgenus Libnotes.
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,500 feet, September 14, 1930 (C. F. Clagg) ; holotype, male.
Limonia (Laosa) manobo is the second species of Laosa to be
described and the first record of the subgenus from the Philip-
pines. It is very different from the subgenotype, gloriosa
46' 3 Alexander: Philippine Tipulidx, XII 449
(Edwards), of French Indo-China, in all details of coloration
and venation, although the beautifully patterned wings are
somewhat alike in the two species. The specific name, manobo,
is that of a native tribe. It should be noted that the very
peculiar structure of the male hypopygium is almost identical
with that of the typical form of Libnotes and that the same
structure has been found in at least one species of the typical
subgenus, Limonia.
LIMONIA (LIMONIA) BILAN sp. nov. Plate 1, fig. 2.
General coloration of mesonotum obscure yellow, the praescu-
tum with three brown stripes; antenna black; flagellar segments
subglobular, with short yellow apical pedicels ; halteres orange ;
legs obscure yellow, the tips of the femora and tibiae darkened ;
wings cream-colored, with a very heavy clouded and spotted
pattern; abdomen dark brown.
Female. — Length, about 11 millimeters; wing, 10.5.
Rostrum and palpi black. Antennae black, the basal flagellar
segments subglobular, with abrupt short yellow apical pedicels ;
penultimate segment short-oval; terminal segment elongate,
pointed at apex, about one-third longer than the penultimate;
verticils longer than the segments. Head dark gray; anterior
vertex (female) a trifle narrower than the diameter of the first
scapal segment.
Pronotum dark brown. Mesonotal prsescutum obscure yel-
low, with three brown stripes, the median stripe broad and
entire, the lateral stripes narrow and becoming subobsolete on
their mesal edges ; scutum with the median area gray, the lobes
chiefly blackened; scutellum large, pale gray; postnotal medio-
tergite blackened. Pleura black, variegated with brown on
the dorsal and ventral sternopleurite and on the meron; dorso-
pleural region restrictedly buffy. Halteres orange. Legs with
the coxae black, the apices restrictedly paler ; trochanters obscure
yellow; femora obscure yellow, the tips deepening to black;
trochanters obscure yellow, the tips narrowly blackened; basi-
tarsi black, the proximal ends brown ; remainder of tarsi black ;
claws (female) with a large outer and two progressively smaller,
more basal spines. Wings (Plate 1, fig. 2) with the very re-
stricted ground cream-colored, the prearcular and costal ground
deeper yellow; a heavy dark brown costal and paler grayish
brown discal pattern; the major brown areas are distributed
along the costa, those at arculus and at the supernumerary cross-
vein in cell Sc more extensive; areas at origin of Rs and end
263774 10
450 The Philippine Journal of Science 1931
of Sc very narrowly divided by a line of the ground color ; stigmal
area in oblique alignment with a band along the cord, crossing
the base of cell R3, the area contiguous with a large spot im-
mediately preceding it; numerous grayish brown spots and
clouds in all the cells, these confluent to form a pattern that is
much more extensive than the ground; veins yellow, darker in
the clouded areas. Venation: Scx ending just before midlength
of Rs, Sc2 at its tip; a weak supernumerary crossvein at near
midlength of cell Sc ; free tip of Sc2 in alignment with R2 ; cell
1st M2 relatively small; m-cu at or just before the fork of M;
anal veins at origin parallel or nearly so.
Abdomen dark brown, the two basal sternites vaguely more
yellowish at base, the succeeding two segments with a linear
yellow median dash ; genital segment obscure fulvous. Oviposi-
tor with the valves reddish horn color ; tergal valves slender and
acute.
Mindanao, Davao district, Mount Apo, altitude 8,000 feet,
September 19, 1930 (C. F. Clagg) ; holotype, female.
Limonia (Limonia) bilan is named from one of the native
tribes living in the vicinity of Mount Apo. It is quite distinct
from the numerous regional species of the subgenus in the abun-
dantly spotted wings, structure of antennae, and details of color-
ation.
LIMONIA (LIMONIA) ATROAURATA sp. nov. Plate 1, fig. 3.
General coloration of head and thorax intense orange, the
mesonotum with two dark brown lines that extend from the
praescutum to the abdomen, converging behind; a narrow black
longitudinal stripe on pleura ; knobs of halteres darkened ; wings
dirty whitish, with a heavy brown clouded and spotted pattern ;
Sc relatively short, Scx ending about opposite one-third the
length of Rs; m-cu at near one-third the length of cell 1st M2.
Female. — Length, about 4.6 millimeters; wing, 5.
Mouth parts very small, black ; palpi reduced, black. Antennae
with the scapal segments black; remainder of organ broken.
Head fiery orange; anterior vertex very broad, at narrowest
point fully three times the diameter of the scape.
Pronotum orange, the anterior notum behind narrowly border-
ed by black. Mesonotal praescutum intense orange, the usual
sublateral stripes represented by brown lines, the broad median
area remaining of the ground color ; extreme lateral margins of
sclerite narrowly and evenly bordered by brownish black, the
46> 3 Alexander: Philippine Tipulidse, XII 451
lines not quite meeting on the cephalic margin ; remaining scle-
rites of mesonotum orange, traversed by narrow brown lines that
converge behind and are direct prolongations of the sublateral
prsescutal stripes, on the postnotal mediotergite strongly ap-
proximated, being divided only by a capillary median line of the
ground color. Pleura orange and yellow, with a narrow black
longitudinal stripe, extending from the cervical sclerites to the
abdomen, the region dorsad of this line more orange, below this
line more yellow ; a linear black streak at the anterior spiracle.
Halteres with the stem obscure yellow, the knobs infuscated.
Legs with the coxae and trochanters yellow; remainder of legs
broken, a single one detached, with the specimen and probably
belonging here, is almost uniformly blackened, the femora a
trifle paler. Wings (Plate 1, fig. 3) dirty whitish, with a heavy
brown pattern consisting of very large clouds and washes; the
major clouds are at arculus; origin of Rs and tip of Sc; stigma;
along cord and outer end of cell 1st M2; beyond midlength of
cells R2 and R3 ; large clouds at ends of anal veins, with an ad-
ditional major area at midlength of cell 2d A; cells R and M
extensively washed with brown ; veins pale, darker in the clouded
areas. Venation: Sc short, Scx ending at near one-third the
length of Rs, Sc2 close to its tip ; Rs relatively short, angulated
and spurred at origin; free tip of Sc2 and R2 in transverse
alignment; cell 1st M2 rectangular, a little shorter than vein
Mi+2 beyond it; m-cu at one-third the length of cell 1st M2,
subequal to the distal section of Cux; anal veins bent rather
strongly into the margin, especially 2d A.
Abdominal tergites velvety black laterally, more brownish
black medially, the caudal margin medially of each segment
with a narrow transverse obscure yellow line, on the basal
tergite much more extensive and almost covering the segment;
subterminal segments more uniformly brown; genital segment
reddish brown; sternites pale brown, the caudal margins nar-
rowly ocherous. Ovipositor with the tergal valves (cerci)
small and strongly upcurved; sternal valves (hypovalvse) longer,
straight, blackened at bases.
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,500 feet, September 14, 1930 (C. F. Clagg) ; holotype, female.
This beautiful little Limonia is very different from any other
fly in the Philippine fauna. The short Sc is distinctive of the
subgenus Limonia; but the distal position of m-cu is a rare con-
dition in this subgenus, being more characteristic of Libnotes.
452 The Philippine Journal of Science i93i
LIMONIA (LIMONIA) BAGOBO sp. nov. Plate 1, fi«. 4; Plate 2, fig. 23.
General coloration obscure yellow; front silvery; antennae,
halteres, and legs blackened; basal flagellar segments subglob-
ular, terminal segment elongate; wings with a blackish tinge;
cell 1st M2 open by atrophy of the basal section of M3; male
hypopygium with the dististyle single, at apex produced into an
acute blackened spine.
Male. — Length, about 3.5 millimeters; wing, 4.2.
Rostrum and palpi very much reduced, black. Antennae black
throughout ; flagellar segments subglobular, the outer ones pass-
ing into oval; terminal segment elongate, narrowed outwardly,
about one-half longer than the penultimate ; verticils short, uni-
laterally arranged, on outer segments becoming smaller and
insignificant. Head brown, the broad frontal region silvery
white.
Mesonotum deep yellow, without distinct markings, the pleura
paler yellow. Halteres dusky, the knobs blackened. Legs with
the coxae and trochanters yeljowish testaceous; remainder of
legs black; claws apparently simple or with setae only. Wings
(Plate 1, fig. 4) with a strong blackish suffusion; veins slightly
darker. Venation : Scx ending about opposite one-third to two-
fifths the length of Rs, Sc2 at its tip ; free tip of Sc2 some distance
before the arcuated R2 ; cell 1st M2 open by the atrophy of the
basal section of M3, cell 2d M2 a trifle longer than its petiole;
m-cu a short distance beyond the fork of M.
Abdomen, including the hypopygium, dark brown. Male hypo-
pygium (Plate 2, fig. 23) with the tergite, 9t, elongate, slightly
longer than wide, the apex bilobed, provided with long conspic-
uous setae. Basistyle, b, elongate, the ventromesal lobe slender.
Dististyle, d, single, oval, narrowed outwardly, at apex produced
into an acute blackened spine; on outer face of basal half with
a circular pale area provided with a small tubercle bearing two
short stout setae. Gonapophyses, g, with the mesal-apical lobe
appearing as an acute blackened hook.
Mindanao, Davao district, Mount Apo, Bakraeyon, altitude
8,000 feet, September 16, 1930 (C. F. Clagg) ; holotype, male.
Limonia (Limonia) bagobo is named from one of the native
tribes inhabiting Mount Apo and surrounding country on the
west side of Davao Gulf. The species is very distinct in the
venation and structure of the male hypopygium. The peculiar
bisetose tubercle on the dististyle of the hypopygium would in-
dicate a relationship with the otherwise very different L. (L.)
canis Alexander and L. (L.) cynotis Alexander.
46» 3 Alexander: Philippine Tipididse, XII 453
LIMONIA (LIMONIA) SUBPACATA sp. nov. Plate 1, fi*. 5; Plate 2, fifr. 24.
Male.— Length about 3 millimeters; wing, 3.8.
Female.— Length, about 3.5 millimeters; wing, 4.
Closely related to L. (L.) pacafo Alexander and L. (L.) pro-
lixicornis Alexander; differing especially in the venation and
structure of the male hypopygium.
Antennas (male) of moderate length, the flagellar segments
short-cylindrical, almost as in subprolixa sp. nov. and much
shorter than in prolixicornis. Head dark.
Thorax light reddish yellow, without distinct markings. Hal-
teres with dusky knobs. Legs chiefly pale testaceous brown,
the outer tarsal segments darkened. Wings (Plate 1, fig. 5)
grayish subhyaline, the stigma not or scarcely differentiated;
veins pale brown. Venation : Sc very short, Sct ending shortly
beyond the origin of Rs, with Sc2 immediately beyond this origin ;
cell 2d A very narrow.
Abdomen reddish brown, the sternites paler. Male hypopyg-
ium (Plate 2, fig. 24) with the lateral lobes of the tergite, 9t,
pale, glabrous, the caudal margin between the lobes emarginate.
Gonapophyses, g, with the lateral lobe darkened, the mesal-apical
lobe pale, very broad, the apex obtusely rounded.
Mindanao, Davao district, Mount Apo, Sibulan River, altitude
7,000 to 8,000 feet, September 21, 1930 (C. F. Clagg) ; holotype,
male; allotype, female.
The present species differs from all described species of the
pacata group in the unusually short Sc which extends only a
short distance beyond the origin of Rs. The male hypopygium
furnishes ready identification characters to separate this fly
from prolixicornis Alexander and subprolixa sp. nov.
LIMONIA (LIMONIA) SUBPROLIXA sp. nov. Plate 1, fig. 6; Plate 2, fiff. 25.
Belongs to the pacata group; antennae of male elongate but
shorter than in prolixicornis; Scx ending beyond midlength of
Rs ; hypopygium with the tergite terminating in two stout lobes,
each bearing five powerful sete; male hypopygium with the
mesal apical lobe of the gonapophyses long and slender.
Male. — Length, about 4 to 4.5 millimeters ; wing, 4.5 to 5.5.
Female. — Length, about 5.5 to 6 millimeters; wing, 5.5 to 5.8.
Rostrum and palpi brownish black. Antennae (male) elong-
ate, but still shorter than in prolixicornis; flagellar segments
cylindrical, with short apical pedicels. Head dark brownish
gray.
Mesonotal prsescutum reddish brown, without distinct mark-
ings, the posterior sclerites of the notum darker medially.
454 The Philippine Journal of Science 1931
Pleura yellow, the dorsal pleurites usually darker. Halteres
dusky, the knobs infuscated. Legs with the fore coxae more or
less darkened on outer face, the other coxae and all trochanters
yellow; remainder of legs brownish black, the femoral bases
restrictedly obscure yellow. Wings (Plate 1, fig. 6) with a
brownish tinge, the oval stigma a trifle darker brown; veins
dark brown. Venation: SCj ending beyond midlength of Rs,
Sc2 a short distance from its tip; cell M2 open by the atrophy
of m ; m-cu at or close to the fork of M.
Abdominal tergites dark brown, the sternites obscure yellow
or brownish yellow. Male hypopygium (Plate 2, fig. 25) with
the tergal plate, 9t, narrow, conspicuous, at apex with two lobes,
each bearing about five stout marginal setae. Basistyles and
dististyles almost as in prolixicornis. Gonapophyses, g, with the
mesal-apical lobe long and slender, gently curved, the apex trun-
cated. j^Edeagus with unusually wide lateral flanges.
Mindanao, Davao district, Mount Apo (C. F. Clagg) ; holo-
type, male, 7,000 to 8,000 feet, September 20, 1930 ; allotype, fe-
male, altitude 8,000 feet, September 19, 1930; paratypes, 15
males and females, 6,500 to 8,000 feet, September 5 to 30, 1930.
Limonia (Limonia) subprolixa is most closely allied to L. (L.)
prolixicornis Alexander, differing in the shorter antennae of the
male and the details of structure of the male hypopygium, espe-
cially the tergite and gonapophyses.
HELIUS (HELIUS) PROCERUS sp. nov. Plate 1, fig. 7; Plate 2, fig. 26.
General coloration dark brown ; rostrum black, slightly longer
than the head; antennae (male) elongate, if bent backward ex-
tending nearly to the base of abdomen ; legs black, the tarsi paling
to yellow ; wings with a faint blackish tinge ; anterior branch of
Rs strongly arcuated at origin and thence running close to and
generally parallel to Rx ; cell 1st M2 long-rectangular, with m-cu
shortly beyond its base.
Male. — Length, about 7 millimeters ; wing, 7.8.
Female. — Length, about 8 millimeters; wing, 7.2.
Rostrum slightly longer than the remainder of head, black;
palpi black. Antennae (male) unusually elongate for this genus,
if bent backward extending nearly to base of abdomen; black
throughout ; flagellar segments cylindrical, with abundant short
dense erect setulae. Antennae (female) short, only a little longer
than the head. Head black.
Pronotum dark medially, obscure yellow laterally. Meso-
notal praeseutum dark brown, without distinct markings; me-
46' 3 Alexander: Philippine Tipulidx, XII 455
dian region of scutum and vicinity of the suture yellow; poste-
rior sclerites of mesonotum darker brown. Pleura dark brown
dorsally, more yellowish brown ventrally. Halteres infuscated.
Legs with the coxae brownish testaceous; trochanters yellowish
testaceous; remainder of legs blackened, the terminal tarsal seg-
ments paling to yellowish. Wings (Plate 1, fig. 7) with a
faint blackish tinge, cells C and Sc dark brown, confluent with
the scarcely differentiated brown stigma; veins dark brown.
Venation: Scx ending some distance beyond r-m, Sc2 faint or
obsolete; r-m on R4+5 shortly beyond the fork of Rs; anterior
branch of Rs very strongly arcuated at base, at the level of the
end of Sc running generally parallel and close to Rx ; Rs nearly
in alignment with the distal section of R4+5; cell 1st M2 long-
rectangular, shorter than any of the veins beyond it; m-cu
a short distance beyond the fork of M.
Abdomen, including the hypopygium, brownish black. Male
hypopygium (Plate 2, fig. 26) with the mesal face of basistyle,
b, at cephalic end with a conspicuous lobe that is covered with
abundant spinous setae. Outer dististyle, od, a simple blackened
rod, the apex entire. Inner dististyle stout and with conspicuous
setae on basal two-thirds, the apex suddenly narrowed. Gona-
pophyses, g, with the mesal angle a long, slender tail-like spine.
Mindanao, Davao district, Mount Apo (C. F. Clagg) ; holo-
type, male, Mainit River, altitude 6,000 feet, September 4, 1930 ;
allotype, female, Galog River, attracted to camp fire, altitude
6,000 feet, September 22, 1930 ; paratype, a fragmentary speci-
men, altitude 7,000 feet, September 11, 1930.
Helms (Helius) procerus is most closely allied to H. (H.)
arcuarius Alexander (Luzon), differing most evidently in the
large size and elongate antennae of the male sex.
HELIUS (HELIUS) APOENSIS sp. nor. Plate 1, fig. 8.
General coloration pale yellow ocherous, without markings;
head blackish gray; wings ocher brown, the stigma a little
darker; wings with cell Rx closed by the apical fusion of veins
R1+2 and R3.
Male. — Length, about 3 millimeters ; wing, 3.5 to 3.6.
Rostrum and palpi black. Antennae black throughout. Head
blackish gray.
Fronotum brown. Mesothorax light yellow ocherous, un-
marked, the scutellum a little paler. Halteres pale, the knobs
slightly darkened. Legs with the coxae and trochanters yellow-
ish testaceous ; remainder of legs pale brownish yellow, the ter-
456 The Philippine Journal of Science 1931
minal tarsal segments brighter yellow. Wings (Plate 1, fig. 8)
pale ocher brown, the pale stigma only slightly indicated; veins
pale brown. Costal fringe (male) conspicuous. Venation : Al-
most as in trianguliferus; anterior branch of Rs shorter and more
erect at origin, the fusion with Rlf2 slightly longer.
Abdomen pale brownish yellow.
Mindanao, Davao district, Mount Apo, altitude 7,000 feet,
September 11, 1930 (C. F. Clagg) ; holotype, male; paratype,
male.
Very similar and closely related to Helius (Helms) triangu-
liferus Alexander (Luzon-Mindanao), differing especially in the
light ocher-yellow coloration of the body.
THAUMASTOPTERA (THAUMASTOPTERA) MACULIVENA sp. nov. Plate 1, fig. 9;
Plate 2, fig. 27.
General coloration pale yellow; antennal scape black, the fla-
gellum yellow; knobs of halteres weakly infuscated; legs pale
yellow, the genua very restrictedly to scarcely darkened; wings
grayish white with a conspicuous brown and gray pattern that
appears as seams to the veins ; Sc relatively short ; r-m shortened
by approximation of the adjoining veins ; male hypopygium with
the dististyle slender, its tip pointed.
Male. — Length, about 2.5 millimeters ; wing, 3.5.
Rostrum brownish black; palpi black. Antennae with the
scape black, the flagellum abruptly light yellow; flagellar seg-
ments subglobular to short-oval, with long conspicuous verticils
that much exceed the segments. Head brown.
Mesonotum pale yellow, in cases the postnotal mediotergite a
trifle darker. Pleura pale yellow. Halteres pale, the knobs
weakly infuscated. Legs with the coxae and trochanters pale yel-
low ; remainder of legs pale yellowish white, the genua very re-
strictedly to almost insensibly darkened. Wings (Plate 1, fig. 9)
with the ground color grayish white, the prearcular and costal
regions clearer cream yellow ; a restricted brown and gray pat-
tern appearing as seams along the veins, arranged as follows:
Arculus, including the surrounding veins ; origin of Rs and op-
posite portion of costa; cord; ends of longitudinal veins from
M1+2 to anal veins, inclusive; a cloud on costa at near three-
fourths the length of cell R2; at midlength of vein R4+5; ni and
adjoining parts of M1+2 and M3; m-cu; at near midlength of
basal section of Cux; a second dash on vein 2d A on basal half; a
weak axillary darkening ; veins pale yellow, brown in the clouded
areas. Costal fringe relatively long. Venation: Sc of mod-
46> 3 Alexander: Philippine Tipididx, XII 457
erate length, Scx ending about opposite one-third the length of
Rs, Sc2 some distance from its tip, opposite or close to origin
of Rs, the latter angulated and long- or short-spurred at origin;
r-m short, reduced by approximation of adjoining veins.
Abdomen yellow, including the hypopygium. Male hypopy-
gium (Plate 2, fig. 27) with the dististyle, d, slender, pale, termi-
nating in an acute pale spinous point, with one long pale seta on
outer margin before apex, together with a row of four black
setae on inner margin, distributed over the outer half; additional
setae on inner face at base. iEdeagus, a, short.
Mindanao, Davao district, Mount Apo, Galog River, altitude
6,000 feet, September 26, 1930; Mainit River, altitude 6,000 to
6,500 feet, September 6 to 14, 1930 (C. F. Clagg) ; holotype,
male; paratypes, 3 males.
It should be noted that this is the first record of the typical
subgenus of Thaumastoptera in the eastern Asiatic area, the only
other member of the genus so far discovered in Asia being Thau-
mastoptera (Taiwanita) issikiana Alexander, from the high
mountains of Formosa. The present species is very distinct from
the genotype, calceatu Mik, in the wing pattern.
HEXATOMINI
ADELPHOMYIA APOANA sp. nov. Plate 1, fig. 10.
General coloration dark brown; antennae 16-segmented, dark
throughout; wings with a faint brown tinge, with a restricted
darker brown pattern, including the stigma and narrow seams
at origin of Rs and along cord; macrotrichia of membrane
relatively sparse.
Female. — Length, about 4 millimeters; wing, 4.3.
Rostrum and palpi black. Antennae black, the flagellar seg-
ments somewhat paler ; sixteen distinct segments, the basal ones
shorter and more crowded ; outer segments long-cylindrical, with
long verticils that exceed the segments ; terminal segment about
one-half longer than the penultimate. Head dark brown.
Thorax almost uniform brown, the central portion of the
praescutum darker. Pleura a trifle more testaceous brown than
the notum. Halteres elongate, dusky, the base of the stem re-
strictedly pale. Legs with the coxae and trochanters yellowish
testaceous; remainder of legs brown, with long outspreading
setae. Wings (Plate 1, fig. 10) with a faint brown tinge, with
a very restricted, slightly darker brown pattern, including the
stigma and narrow seams at origin of Rs and along the cord;
458 The Philippine Journal of Science 1931
veins pale brown. Macrotrichia of cells relatively sparse, in
the outer ends of cells R2 to Ms, inclusive. Venation : Scx ending
shortly before the fork of Rs, Sc2 some distance from its tip;
Rs weakly angulated at origin ; m-cu at near midlength of lower
face of cell 1st M2 ; cell Mx present.
Abdomen brownish black. Ovipositor with the elongate
tergal valves darkened at bases, the slightly upcurved acute
tips yellow.
Mindanao, Davao district, Mount Apo, Kidopawan trail to
Lake Lino, altitude 7,000 to 8,000 feet, September 20, 1930
(C. F. Clagg) ; holotype, female.
Adelphomyia apoana is apparently distinct from any of the
now rather numerous regional species in the wing pattern,
venation, and conformation, and in the relatively sparse macro-
trichia of the membrane. The nearest ally seems to be A.
carbonicolor Alexander.
ADELPHOMYIA PAUCISETOSA sp. nov. Plate 1, fig. 11; Plate 2, fig. 28.
General coloration black; antennae 15-segmented, the fusion
segment yellow, remainder of organ darkened ; wings milk white
with a heavy brown pattern that is distributed chiefly as narrow
broken crossbands ; macrotrichia of membrane very sparse, being
restricted to a few trichia in ends of cells R3 and R4 ; male hypo-
pygium with the outer dististyle bearing a long erect spine on
inner face at near midlength.
Male. — Length, about 3 millimeters ; wing, 3.8.
Rostrum and palpi brownish black. Antennae with the scape
black, the fusion segment pale yellow; remainder of flagellum
brown ; antennae with fifteen segments, the short-conical fusion
segment involving two segments; outer flagellar segments sub-
cylindrical, with verticils that exceed the segments in length;
terminal segment about one-fourth longer than the penultimate.
Head black.
Pronotum obscure brownish yellow medially, blackened later-
ally. Mesonotal praescutum yellowish brown to chestnut, darker
medially; scutal lobes light brown; posterior sclerites of meso-
notum dark brown. Pleura black. Halteres chiefly pale yel-
low, the central portion of stem vaguely darker. Legs with the
fore coxae brownish yellow, the remaining coxae black ; trochan-
ters testaceous; remainder of legs brown, the outer tarsal seg-
ments somewhat darker ; no tibial spurs ; segments of legs with
long conspicuous setae. Wings (Plate 1, fig. 11) milky white,
with a heavy brown pattern that is arranged chiefly as six or
46, 3 Alexander: Philippine Tipulidm, XII 459
seven, narrow, broken crossbands, interrupted at the central
portion of the disk ; basal band beyond arculus, complete ; second
band at origin of Rs and end of vein 2d A, broken in cells M and
Cu ; third band at Sc2 and end of 1st A, interrupted but replaced
in a slightly more distal position by a similar seam along cord ; an
interrupted irregular band includes the stigma, outer end of
cells 1st M2 and M4; an outer band includes end of R3, and a
prolongation of the area across cells R5 and 2d M2; additional
brown clouds at ends of veins R4 and R5; paler washes in cells
M, Cu, and at midlength of cell 2d A ; veins pale, darker in the
clouded areas. Macrotrichia of cells very sparse, being restrict-
ed to a group of five or six in outer end of cell R4, with one or
two more in cell R3. Venation: Scx ending about opposite the
end of Rs ; veins R3 and R4 slightly upcurved at ends ; R2 at fork
of R3+4; cell Mx present; cell 1st M2 strongly narrowed at proxi-
mal end, r-m being correspondingly lengthened, arcuated.
Abdomen chiefly black, including the hypopygium. Male
hypopygium (Plate 2, fig. 28) with the outer dististyle, od, an
elongate-oval blackened structure, terminating in two slender
spines, one being slightly more curved; just beyond midlength
of style on inner margin a long slender erect spine. Inner dis-
tistyle, id, very stout at base, the obtuse tip narrowed.
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,000 feet, September 22, 1930 (C. F. Clagg) ; holotype, male.
Adelphomyia paucisetosa most closely resembles A. nebulosa
(de Meijere), of western Java, differing from all known species
in the very notable reduction in number of macrotrichia of the
wings, a condition which presages their total loss.
EPIPHRAGMA (POLYPHRAGMA) FUSCOFASCIATA gp. nov. Plate 1, fig. 12.
General coloration of mesonotum ocherous brown, dark brown
laterally; pleura and pleurotergite black; antennal scape and
fusion segment of antenna pale; halteres black; wings yellow,
with three more or less complete crossbands of brownish black,
the third band at the cord, very broad but more or less interrupt-
ed by pale ; wing tip pale, with small dark spots at ends of the
veins.
Female. — Length, about 7 millimeters; wing, 7.5.
Rostrum yellowish gray; palpi black. Antennae with the first
scapal segment light brown; second segment obscure yellow;
fusion segment bright orange; remainder of flagellum black.
Head yellowish gray, the central and posterior portions of the
vertex darker.
460 The Philippine Journal of Science lost
Pronotum obscure yellow, deepening to black on sides. Meso-
notal praescutum ocher brown sublaterally, darker brown medial-
ly, the lateral margins narrowly and abruptly dark brown ; scutal
lobes dark brown; scutellum black, the parascutella somewhat
paler ; postnotal mediotergite obscure yellowish brown, blackened
posteriorly. Pleura and pleurotergite black. Halteres black.
Legs with the fore coxse dark brown, the remaining coxae black ;
trochanters brownish yellow; femora yellow, darkened subter-
minally; remainder of legs yellow. Wings (Plate 1, fig. 12)
with the ground color yellow, with three heavy crossbands of
brown to brownish black; basal area including the arcular re-
gion; second band at origin of Rs; third band very broad, ex-
tending from before the cord to the level of R3, interrupted by a
few small yellow areas, as in cells Sc2, R2, 1st M2, M3, and M4;
wing apex pale, varied by a series of marginal brown areas at
ends of veins R4 to M2, inclusive; the yellow alternating cross-
bands are slightly clouded with dusky in the cubital and anal
fields, leaving clear yellow margins bordering the crossbands;
veins yellow, dark in the infuscated areas. In the paratypes, the
outer band is more extensively interrupted by pale markings.
Venation: Crossveins and spurs in cell C very much restricted
in number ; Rs square and weakly spurred at origin ; cell 1st M2
relatively small.
Abdomen rather light brown, the caudal margins of the seg-
ments narrowly but conspicuously brownish black; genital seg-
ment obscure yellow ; valves of ovipositor horn-colored, the bases
of the cerci darker.
Mindanao, Davao district, Mount Apo (C. F. Clagg) ; holotype,
female, Galog River, altitude 5,000 to 6,000 feet, September 12,
1930; paratypes, two females, Sibulan River, altitude 7,000 to
8,000 feet, September 21, 1930; one female, Kidopawan trail
from Lake Lino, altitude 7,000 to 8,000 feet, September 20, 1930.
Epiphragma (Polyphragma) fuscofasciata is distinguished
from other members of the ochrinota group by the handsomely
banded wing pattern.
EPIPHRAGMA (POLYPHRAGMA) LATITERGATA sp. nov. Plate 2, fiff. 29.
General coloration of mesonotum brownish yellow, contrasting
with the blackened pleura ; legs yellow, the femora with a broad
pale yellow subterminal ring; wings with the ground color light
brown, with a heavier brown pattern that is narrowly margined
with light yellow; male hypopygium with the lateral lobes of
the tergite broad, obtuse; interbasal process at apex expanded
46' s Alexander: Philippine Tipulidse, XII 461
at apex into a truncated blade, the outer apical angle bearing a
small, curved, beaklike spine.
Male. — Length, about 7.5 to 8 millimeters ; wing, 8 to 9.
Female.— Length, about 9 to 10 millimeters ; wing, 9 to 9.5.
Rostrum and palpi dark brown. Antennae with the first scapal
segment blackened, the second obscure brownish yellow; basal
flagellar segments not distinctly united into a fusion segment,
beyond the base black, the verticils exceeding the segments in
length. Head dull yellowish gray, the posterior vertex more
reddish brown, the caudal portions more infuscated on either side
of the midline.
Mesonotum dull brownish yellow, without markings, the lateral
portions of the praescutum deep chestnut orange. Pleura black-
ened, as in the group, the ventral sternopleurite remaining yellow-
ish. Halteres yellow, the knobs dark brown. Legs with the
coxae obscure yellow, narrowly darkened basally, especially the
posterior coxae ; trochanters yellow ; femora yellow, with a broad
pale brown subterminal ring; remainder of legs light yellow,
the terminal tarsal segments passing into fulvous. Wings with
the ground color light brown with a heavier brown pattern,
arranged as in the group, the major areas being at arculus;
origin of Rs ; along cord and outer end of cell 1st M2 ; fork of
Mi +2 ; and as conspicuous circular marginal clouds at ends of
all longitudinal veins ; the dark pattern is narrowly but conspic-
uously bordered by pale yellow; costal margin yellow, beyond
the region of the stigma appearing as three isolated spots in
outer ends of cells R2, R3, and R4; veins dark, obscure yellow in
the costal interspaces. No dilation of the axillary region. Ve-
nation: Spurs and supernumerary crossveins in cell C six to
eight in number, all seamed by darker ; Rs relatively long, angu-
lated and weakly spurred at origin; m-cu variable in position,
at one-fourth to midlength of cell 1st M2.
Abdomen chiefly dark brown, including the sternites and hypo-
pygium. Male hypopygium (Plate 2, fig. 29) generally as in
fulvinota but differing in some important regards, notably the
broad, obtuse lobes of the tergite, 9£, and the shape of the inter-
basal processes, i. These latter normally are expanded at apex
into a squarely truncated blade that bears on outer apical angle
a small, curved hooklike spine.
Mindanao, Davao district, Mount Apo, altitude 5,000 to 8,000
feet, August 31 to September 21, 1930 (C. F. Clagg) ; holotype,
male; allotype, female; paratopotypes, several of both sexes.
462 The Philippine Journal of Science mi
Among the species of the ochrinota group, the present fly is
closest to Epiphragma {Polyphragma) fvlvinota Alexander,
from which it differs most evidently in the wing pattern, with
conspicuous narrow yellow margins to the darkened areas, the
paler brown femoral annuli, and the structure of the male hypo-
pygium, notably of the tergite and interbasal processes.
EPIPHRAGMA (POLYPHRAGMA) NIGROTIBIATA sp. nov. Plate 1, fig. 13; Plate 2,
fig. 30.
General coloration of mesonotum yellow, variegated with dark
brown; pleura yellow, with scattered small dark brown spots;
femora yellow basally, the distal half black, inclosing two narrow
yellow rings ; tibiae black ; tarsi yellow ; wings brownish yellow,
the cephalic portion deeper yellow, the surface with a heavy
brown pattern.
Male. — Length, about 7.5 millimeters; wing, 8.5.
Rostrum and palpi black. Antennae relatively short; scapal
segments brown, the fusion segment and second segment of
flagellum orange ; remainder of flagellum black ; verticils exceed-
ing the segments in length. Head brownish gray, the lateral
portions of the vertex and the genae more reddish brown.
Pronotum yellow, the anterior notum variegated with dark
brown on the sides. Mesonotal praescutum yellow, variegated
with dark brown medially, the area broad and entire behind, be-
coming bifid and obsolete in front; sublateral portions of the
sclerite deeper reddish yellow than the pollinose interspaces;
extreme lateral margins of praescutum dark brown ; scutal lobes
reddish brown, margined with slightly darker brown, the ce-
phalic lateral portions brighter ; scutellum brown ; postnotal me-
diotergite dark brown, pruinose. Pleura yellow, variegated with
scattered brown areas, located on the dorsal anepisternum, dorsal
sternopleurite, ventral sternopleurite, meron, and dorsal and
ventral pleurotergite. Halteres dark brown, the base of the stem
light yellow. Legs with the coxae and trochanters yellow, the pos-
terior coxae a little darker apically; femora yellow basally, the
outer half passing into black, inclosing a narrow apical and a
slightly wider subapical yellow ring; tibiae black, the extreme
base yellow ; tarsi light yellow, the terminal segments darkened.
Wings (Plate 1, fig. 13) brownish yellow, the prearcular, costal,
and radial fields deeper yellow; a heavy brown pattern, distri-
buted as follows : A series of narrow costal and subcostal areas
surrounding the crossveins and spurs in the former cell ; larger
areas at arculus ; origin of Rs ; along cord ; outer end of cell 1st
46' 3 Alexander: Philippine Tipulidse, XII 463
M;; fork of M1+2 ; marginal clouds at ends of all longitudinal
veins, largest on the anals; a restricted dark area in axillary re-
gion; radial and medial cells beyond the level of the fork of M1+2
extensively darkened, confluent with the marginal dark areas
in this field to produce a radiate appearance; dark areas behind
the costa narrowly bordered by cream yellow; veins pale brown,
darker in the infuscated areas. Venation: A series of super-
numerary crossveins and spurs in cell C ; m-cu more than one-
half its length beyond the fork of M ; supernumerary crossvein
in cell Cu atrophied or nearly so.
Abdominal tergites dark brown, the basal ring of the second
segment obscure yellow laterally ; impressed transverse lines of
the remaining tergites narrowly bordered by pale; sternites
obscure yellow, the extreme caudal margins of the segments
darkened. Male hypopygium (Plate 2, fig. 30) with the inter-
basal process, i, a slender rod from a dilated base, the apex
weakly expanded and further produced into a small curved point.
Outer dististyle, od, dilated at midlength, the apex a strongly
curved spine.
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,500 feet, September 14, 1930 (C. F. Clagg) ; holotype, male.
Epiphragma (Polyphragma) nigrotibiata is well-distinguished
by the uniformly black tibiae and the pattern of the femora.
EPIPHRAGMA (POLYPHRAGMA) APOENSIS sp. nov. Plate 1, &g. 14; Plate 2, fig. 31.
General coloration of mesonotum yellow, the disk with three
confluent brown stripes; pleura chiefly yellow, margined with
brownish black; femora yellow with a broad black subterminal
ring; wings pale brown, with a heavy dark brown pattern that
is narrowly bordered by cream yellow; male hypopygium with
the lateral lobes of the tergite broad ; interbasal process a simple
blade terminating in a small beak.
Male. — Length, about 7.5 millimeters; wing, 8.5.
Rostrum brown; palpi black. Antennae with the scape light
brown; fusion segment small, yellow; remainder of flagellum
black; basal flagellar segments short-oval, the outer segments
subcylindrical, with verticils that are about as long as the seg-
ments. Head above with the central area dark brown, paling
to reddish on sides of posterior vertex.
Pronotum yellow, dark brown laterally. Mesonotal prsescu-
tum yellow, the extreme lateral margin dark brown; disk of
prsescutum almost covered by three confluent brown stripes that
are further divided by a capillary dark brown vitta ; scutal lobes
464 The Philippine Journal of Science 1931
brown, the extreme cephalic-lateral angles brightened ; posterior
sclerites of mesonotum yellowish brown, the postnotal medio-
tergite darker medially. Pleura chiefly yellow, variegated with
brownish black on the margins, including the dorsopleural mem-
brane, cephalic and ventral margin of sternopleurite, meron and
dorsal and ventral portions of pleurotergite. Halteres long,
pale yellow, the knobs infuscated. Legs with the coxae and tro-
chanters yellow; only a single (hind) leg remains; femora yel-
low, brighter yellow on distal fourth, this area inclosing a broad
black ring; tibiae and tarsi yellow. Wings (Plate 1, fig. 14)
with the ground color pale brown, with a heavy dark brown
pattern that is bordered by narrow cream-yellow margins ; costal
brown pattern including both cells C and Sc, with three costal
areas passing into a large solid marking at origin of Rs; an
hourglass-shaped darkening at the cord ; wing apex beyond cell
1st M2 chiefly darkened, variegated by yellow marginal areas
in the outer ends of cells R3, R4, Mly and 2d M2, together with
small paler yellow spots in bases of cells M„ 2d M2, and M3 ; a
large darkened mark at end of vein 2d A, extending to Cu ; axilla
darkened; a large area at arculus; veins pale yellow in the
ground, darker in the clouded portions. Venation : Costal spurs
and crossveins numerous, including about four beyond the origin
of Rs, the latter angulated and spurred at origin; m-cu about
one-half its length beyond the fork of M.
Abdominal tergites dark brown, the basal ring brighter, espe-
cially laterally; sternites extensively yellowish, the caudal mar-
gins darkened ; hypopygium chiefly darkened. Male hypopygium
(Plate 2, fig. 31) with the lateral lobes of the ninth tergite, 94,
broad, separated by a deep notch. Interbasal process, i, a rela-
tively narrow blade, the apex a small curved beak. Outer dis-
tistyle, od, with the main body spinous on outer margin, the apex
a long curved spine.
Mindanao, Davao district, Mount Apo, Seliban River, altitude
7,000 feet, September 11, 1930 (C. F. Clagg) ; holotype, male.
Belongs to the fuscosternata group, having the mesonotum and
pleura conspicuously variegated yellow and brown. The type of
hypopygium is much like that of E. (P.) fvlvinota that belongs
to the ochrinota group, the resemblance being especially striking
in the general features of the interbasal process and dististyles.
EPIPHRAGMA (POLYPHRAGMA) HASTATA sp. nov. Plate 2, fig. 32.
General coloration of mesonotal prsescutum dark brown,
margined with yellow; pleura yellow, variegated with dark
brown ; femora yellow, with a broad subterminal dark brown to
46, 3 Alexander: Philippine Tipulidx, XII 465
brownish black ring; wings with a heavy dark brown pattern
that is bordered by cream yellow; male hypopygium with the
lobes of the ninth tergite broad, microscopically roughened at
apices ; interbasal rod an acute spearlike point.
Male. — Length, about 9 millimeters ; wing, 10.
Rostrum light brown ; palpi dark brown. Antennae with the
scape brownish yellow; basal three flagellar segments light
yellow, the remainder passing into dark brown; no distinctly
developed fusion segment. Head orange, the center of the ver-
tex infuscated.
Pronotum yellow. Mesonotal prsescutum yellow laterally,
margined narrowly with dark brown; disk almost covered by
three confluent dark brown stripes, the region of the interspaces
more yellowish pollinose ; scutal lobes dark brown ; median area
of scutum and the scutellum pale, yellowish pollinose ; postnotal
mediotergite brown, with a more yellow pollinose area on either
side at midlength. Pleura yellow pollinose, variegated with
dark brown, including the anterior dorsopleural region, the
anterior margin of the anepisternum and sternopleurite, the
meron, and the dorsal and ventral pleurotergite. Halteres pale
yellow, the knobs infuscated. Legs with the coxae and tro-
chanters orange yellow, the posterior coxae and cephalic face
of the fore coxae darkened; femora yellow, with a very broad
dark brown (fore femora) to brownish black (posterior femora)
subterminal ring; remainder of legs yellow. Wings with the
ground color pale brown, with a heavy dark brown pattern;
prearcular and costal portions deeper yellow; brown areas bor-
dered by creamy margins; dark markings in cells C and Sc
numerous; major dark areas arranged as follows: Arculus; ori-
gin of Rs, with a more-elongate area in alignment at the super-
numerary crossvein in cell Cu and end of vein 2d A, interrupted
at cell M ; along cord, narrowed in the medial field ; outer end of
cell 1st M2 ; ends of all longitudinal veins, continued back along
the veins; veins light brown, darker in the infuscated areas,
more yellow in the flavous interspaces.
Abdominal tergites chiefly dark brown, the basal rings paler;
sternites more yellowish, the incisuress narrowly darkened ; hypo-
pygium with the basistyles pale. Male hypopygium (Plate 2,
fig. 32) with the lateral lobes of the tergite, 9t, broad, micro-
scopically roughened at apices, separated by a deep U-shaped
notch. Interbasal process, i, an acute spearlike rod. Outer dis-
tistyle, od, terminating in an acute curved spine.
263774 11
466 The Philippine Journal of Science 1931
Mindanao, Davao district, Mount Apo (C. F. Clagg) ; holo-
type, male, altitude 6,000 feet, August 30, 1930 ; allotype, female,
altitude 7,000 feet, September 11, 1930.
Epiphragma (Polyphragma) hastata belongs to the fuscoster-
%ata group, being most closely allied to E. (P.) fttscosternata
Alexander and E. (P.) apoensis sp. nov. It differs from the
latter in the distinctive structure of the male hypopygium and
from the former (the male of which is still unknown) in the
more-restricted amount of dark coloring in the anal cells of the
wing.
EPIPHRAGMA (POLYPHRAGMA) CANINOTA sp. nov. Plate 1, &g. 15; Plate 2, fig. 33.
General coloration of dorsum of head and mesonotum light
ashy gray; knobs of halteres infuscated; legs yellow; wings of
both sexes with a conspicuous axillary crenulation; radial cells
clouded with brown; darkened areas of wing not bordered by
paler.
Male. — Length, about 6.5 millimeters; wing, 7.5.
Female. — Length, about 8.5 millimeters ; wing, 8.2.
Rostrum reduced, pale brown. Antennae with the scape and
fusion segment pale yellow, the remainder of the flagellum
black. Head above light ashy gray, the posterior slope of the
vertex, together with the gense, more orange yellow, infuscated
medially.
Mesonotum above light ashy gray on the dorsomedian portion,
the sides of the praescutum and postnotal mediotergite abruptly
orange yellow. Pleura yellow. Halteres yellow, the knobs in-
fuscated. Legs yellow, the terminal tarsal segments darkened.
Wings (Plate 1, fig. 15) yellowish brown, the costal margin light
yellow, continued to the wing tip in the radial field but here
broken into spots by brown clouds at the ends of the veins;
radial field extensively ssuffused with brown; additional brown
clouds and spots at arculus ; origin of Rs ; cord ; outer end of cell
1st M2; fork of Mi+2; at supernumerary crossveins in cells C
and Cu, and as large marginal clouds at ends of the veins ; veins
brownish yellow, darker in the clouded areas. Axillary crenu-
lation large and conspicuous, a trifle less developed in female
than in male. Venation: Supernumerary crossvein in cell Cu
well-preserved in both sexes ; m-cu in male at fork of M, in fe-
male, beyond the fork but with the crossvein in transverse align-
ment with the other elements of the cord.
Abdominal tergites yellowish brown, darker laterally; stern-
ites clearer yellow. Male hypopygium (Plate 2, fig. 33) with
46, 3 Alexander: Philippine Tipulidx, XII 467
the apex of the interbasal process, i, a tonglike structure, the
lateral arm being a curved spine. Outer dististyle relatively
slender, the vestiture of outer face consisting of abundant del-
icate setulse, with a few longer setae. Inner dististyle with apex
dilated into a slight head, bearing one unusually long seta.
Mindanao, Davao district, Mount Apo, Galog River, altitude
6,000 feet, September 8, 1930 (C. F. Clagg) ; holotype, male;
allotype, female, in copula.
There is a considerable group of species of Polijphragma in
the Philippines having the head and mesonotum chiefly clear ashy
gray, differing from one another by distinctions in the degree of
development of the axillary lobe, the wing pattern, and slight
details of structure of the male hypopygium. I have called this
group of flies the crenulata group. The present fly falls in this
division and seems closest to E. (P.) cinereinota Alexander;
which differs in the coloration of wing and body, as the blackened
subterminal ring of the abdomen.
EPIPHRAGMA (POLYPHRAGMA) GRISEICAPILLA sp. nov. Plate 1, fig. 16; Plate 2,
figr. 34.
Belongs to the crenulata group ; general coloration of dorsum
of head and mesonotum light ashy gray; antennal scape dark
brown, the flagellar fusion segment light yellow ; wings with the
ground color brownish yellow, the costal region clearer yellow;
a heavy brown pattern that is narrowly bordered by clear yel-
low; male hypopygium with the apex of the interbasal process
expanded, the notch small, the lobes broadly flattened.
Male. — Length, about 7 millimeters; wing, 7.5.
Rostrum and palpi black. Antennae with the scapal segments
dark brown, sparsely pruinose ; fusion segment yellow ; remain-
der of flagellum black ; verticils longer than the segments. Head
light gray in front, behind and on sides more brownish, the
center of the posterior vertex brownish black.
Mesonotum clear light gray, the suture medially more bright-
ened; lateral portions of the prcescutum broadly and abruptly
orange yellow. Pleura yellow. Halteres obscure yellow, the
knobs inf uscated. Legs with the coxae and trochanters yellow ;
remainder of legs yellow, the femora a trifle darker just before
the tips, this coloration caused more especially by an increase
in dark setae; terminal tarsal segments only slightly darkened.
Wings (Plate 1, fig. 16) with the ground color brownish yellow,
the cells beyond the cord even more suffused; prearcular and
costal regions clear yellow, beyond the end of Sc continued to
468 The Philippine Journal of Science 1931
the wing tip as yellow spots in the outer ends of cells R2, R3, and
R4 ; darker brown areas at arculus ; origin of Rs ; cord ; fork of
R2+3-f4; outer end of cell 1st M2; fork of Mlf2; supernumerary
crossvein in cell Cu, and the marginal clouds, all these areas
narrowly bordered by clearer yellow rings; veins dark brown,
darker in the clouded areas. Axillary crenulation of moderate
size only, about one-half as deep as in the corresponding sex
of crenulata or caninota. No macrotrichia on Rs or R2+3+4.
Venation : m-cu nearly its own length beyond the fork of M.
Abdominal tergites yellowish brown, the sternites clearer yel-
low, with the incisures narrowly darkened ; hypopygium brown-
ish yellow. Male hypopygium (Plate 2, fig. 34) much as in
caninota, but the interbasal process, i, differently constructed,
the apical notch being very small and shallow, the lobes broadly
flattened.
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,000 feet, September 16, 1930 (C. F. Clagg) ; holotype, male.
Epiphragma (Polyphragma) griseicapilla is allied to E. (P.)
crenulata Alexander and E. (P.) caninota sp. nov., in the general
coloration and relatively deep crenulation of the wing axilla,
differing in the wing pattern and details of structure of the hypo-
pygium.
EPIPHRAGMA (POLYPHRAGMA) ANGUSTICRENULA sp. nov. Plate 1, fi*. 17; Plate
2, fig. 35.
Belongs to the crenulata group ; general coloration of head and
mesonotum light ashy gray; wings with a yellowish brown
ground color, the dark pattern but slightly evident against this
ground and not margined with paler; axillary crenulation of
wing very shallow ; male hypopygium with the interbasal process
bifid at tip, the lateral arm a slender curved spine.
Male. — Length, about 7.5 millimeters; wing, 8.2.
Rostrum and palpi black. Antennae with the scape and fusion
segment obscure brownish yellow ; remainder of flagellum black ;
fusion segment oval, involving three segments ; verticils of flagel-
lum exceeding the segments in length. Dorsum of head on
front and anterior vertex light gray, the posterior vertex dark
reddish brown, more blackened medially.
Mesonotum above light gray, the lateral margins of the prae-
scutum abruptly orange yellow. Pleura obscure yellow, the dor-
sopleural region slightly darkened. Halteres dusky, the knobs
infuscated. Legs with the coxae and trochanters yellow; re-
mainder of legs yellow, the terminal tarsal segments darkened.
46, 3 Alexander: Philippine Tipulidx, XII 469
Wings (Plate 1, fig. 17) with a yellowish brown suffusion, the
preareular and costal regions more yellowish, variegated by
brown clouds at the veins; disk of wing with a diffuse brown
pattern that is little conspicuous against the ground color, the
areas not bordered by brighter; veins brown, yellow in the
flavous costal interspaces. Axillary crenulation very shallow
for this group of the subgenus, being about as wide as the prear-
eular cell immediately cephalad of it. Venation: Costal cross-
veins and spurs few, but strong and complete ; m-cu about one-
half its length beyond the fork of M.
Abdominal tergites light brown, bordered by dark brown later-
ally, the sternites yellow, with narrow darker margins. Male
hypopygium (Plate 2, fig. 35) with the interbasal rods, i, bifid
at tips, the lateral arm a slender curved spine, much as in
crenulata, the mesal arm short and broadly truncated. Outer
dististyle, od, relatively slender, the tip a chitinized, gently curved
spine.
Mindanao, Davao district, Mount Apo, Kidapawan trail to
Lino Lake, altitude 7,000 to 8,000 feet, September 20, 1930 (C. F.
Clagg) ; holotype, male.
Epiphragma (Polyphragma) angttsticrenula differs from the
other species of this group of the subgenus in the scarcely de-
veloped axillary crenulation of the wing, in conjunction with the
other characters listed above.
ERIOPTERINI
TRENTEPOHLIA (PARAMONGOMA) CHIONOPODA sp. nov.
General coloration of thorax yellow; tips of femora white;
tibiae and tarsi white, the basal half of the former more-obscure
whitish ; wings grayish subhyaline, the preareular and costal re-
gions more yellowish.
Male. — Length, about 4 millimeters; wing, 4.2.
Rostrum and palpi brown. Antennae with the scape dark
brown, the flagellum somewhat lighter in color; flagellar verti-
cils a little longer than the segments.
Thorax uniformly yellow. Halteres pale, the knobs weakly
dusky. Legs with the coxae and trochanters yellow; femora
dirty white, the tips paling to clear white ; tibiae and tarsi white,
the basal half of the former a trifle more obscure. Wings gray-
ish subhyaline, the preareular and costal regions light yellow;
stigma small and very vague ; veins pale brown, Sc light yellow.
Venation: R2 close to fork of R3+4; R3 less perpendicular and
cell 1st M2 smaller than in banahaoensis; cell 2d M2 narrow.
470 The Philippine Journal of Science 1931
Abdominal tergites brown medially, paler laterally; sternites
light yellow, the outer segments more infuscated; hypopygium
yellow.
Mindanao, Davao district, Mount Apo, Galog River, altitude
6,000 feet, at trap lantern, September 13, 1930 (C. F. Clagg) ;
holotype, male.
Trentepohlia (Paramongoma) chionopoda is readily told from
the other regional species by the coloration of the legs. The
type of the subgenus Paramongoma, albitarsis (Doleschall), of
Amboina, still seems to be known only from DoleschalPs insuffi-
cient description and faulty figure, which, if only approximately
correct, serve to separate the two species of crane flies.
TRENTEPOHLIA (PARAMONGOMA) PUSILLA Edwards.
Trentepohlia (Paramongoma) pusilla Edwards, Treubia 9 (1927)
356.
Mindanao, Davao district, Lawa, at light, April, 1930 (C. F.
Clagg). This species was described from Sebesi Island, near
Krakatau, Java, where it was taken in April, 1921, by Dammer-
man.
The present specimen agrees almost exactly with Edwards's
description. The allied T. (P.) banahaoensis Alexander (Luzon)
has R3 short and more nearly erect and the tips of the femora
narrowly but conspicuously whitened.
TRENTEPOHLIA (MONGOMA) ^QUIALBA sp. nov. Plate 1, fig. 18.
General coloration of mesonotum orange fulvous, patterned
with black; femora light brown, the tips abruptly snowy white,
the amount subequal on all legs ; bases and tips of tibiae whitened ;
wings with cells C and Sc strongly blackened, the prearcular
region pale ; abdominal tergites yellow, with a broad black dorso-
median stripe.
Male. — Length, about 14 to 16 millimeters; wing, 8.2 to 8.6.
Female. — Length, about 14 millimeters ; wing, 9.
Rostrum and labial palpi obscure yellow ; maxillary palpi black.
Antennae with the scapal segments brown, the flagellum black;
flagellar segments long-cylindrical, with verticils that are sub-
equal to the segments. Head fulvous orange, the vertex carinate
medially.
Mesonotal praeseutum orange fulvous, narrowly darkened
laterally ; centers of scutal lobes darkened ; scutellum testaceous
brown, darker brown caudally; postnotal mediotergite black
46, 3 Alexander: Philippine Tipulidse, XII 471
posteriorly and on sides, a little paler medially in front. Pleura
chiefly orange yellow, the ventral pleurites slightly white prui-
nose; in cases the pleurotergite a little darkened posteriorly.
Halteres blackened, the base of the stem restrictedly yellow.
Legs with the coxae and trochanters orange yellow; femora
light brown, darkened outwardly, the tips abruptly snowy white,
the amount subequal on all legs; tibise dark brown, the bases
narrowly, the tips more broadly whitened ; tarsi white, the ter-
minal segment a little darkened; femora with short black setae
distributed over the entire length, with two or three longer black
setae at apex. Wings (Plate 1, fig. 18) narrow, cells C and Sc
strongly blackened, confluent with the slightly darker stigma;
wing apex narrowly infuscated; vague and narrow, scarcely
evident dark seams on posterior cord ; cell Cu and a spot between
anal veins at point of divergence dark brown ; prearcular region
pale; veins black, the outer branches of M paler. Venation:
R2 about two-thirds to three-fourths its length before fork of
R3+4; veins R3 and R4 very strongly divergent; m-cu at or
shortly before fork of M; fusion of C\x1 and 1st A very slight
to punctiform.
Abdominal tergites yellow laterally, with a broad black dorso-
median stripe ; sternites more extensively yellow, the subcaudal
margins narrowly darkened ; extreme apices of segments silvery ;
subterminal segments and hypopygium blackened.
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,500 feet, September 14, 1930 ; Galog River trail, altitude 5,000
to 6,000 feet, September 12, 1930 (C. F. Clagg) ; holotype, male;
allotype, female; paratypes, 2 males.
The present species, and the two next defined, are all allied
to T. (M.) luzonensis Edwards, from which they may be separat-
ed by the following key :
1. Tibiae of all legs approximately alike in color 2.
Tips of fore tibiae broadly blackened, of posterior tibiae even more broadly
snowy white T. (M.) luzonensis Edwards.
2. Tips of femora and tibiae snowy white T. (M.) szquwlba sp. nov.
Tips of femora and tibiae blackened 3.
3. Mesonotal praescutum polished black, the humeral region yellow; a dark
area on anepisternum ; abdominal tergites blackened.
T. (M.) s&quinigra sp. nov.
Mesonotal praescutum yellow, in cases restrictedly darkened at suture;
pleura uniformly pale; abdominal tergites yellow with a narrow, more
or less interrupted, dorsomedian black stripe.
T. (M.) maju!&cula sp. nov.
472 The Philippine Journal of Science mi
TRENTEPOHLIA (MONOOMA) iEQUINIGRA sp. nov. Plate 1, figr. 19.
General coloration of mesonotum polished black, the humeral
region of the praescutum extensively yellow; pleura yellow, the
dorsal anepisternum darkened; femora yellow, the tips of all
narrowly but conspicuously blackened, the amount subequal on
all legs; fore femora (male) broadly darkened on central portion;
wings narrow, whitish, the costal border light yellow; wing
tip narrowly darkened; abdominal tergites and a subterminal
ring black, the sternites light yellow.
Male. — Length, about 13 millimeters; wing, 8.5 by 1.6.
Female. — Length, about 10 to 13 millimeters ; wing, 7.2 by 1.5
to 9 by 1.75.
Rostrum and palpi brownish black. Antennae with the scapal
segments black ; flagellum broken. Head brownish gray, clearer
gray in front, the vertex carinate.
Pronotum obscure yellow. Mesonotal praescutum polished yel-
low, the lateral margins as far cephalad as the pseudosutural
f ovese, together with a median line almost to the cephalic margin,
blackened, leaving the humeral region extensively of the ground
color; posterior sclerites of mesonotum chiefly blackened, the
median area of the scutum a little brighter. Pleura abruptly
yellow, with a large dark area on the dorsal anepisternum.
Halter es brownish black, the base of the stem brightened. Legs
with the coxae and trochanters yellow; femora yellow, the tips
of all legs somewhat narrowly but conspicuously blackened, the
amount equal on all legs ; in male, the general coloration of the
fore femora is darker brown in the central portion, the tips
again dark brown as described; tibiae obscure yellow, the tipb
blackened; tarsi yellow; all femora with small scattered black
setae distributed over the entire length. Wings (Plate 1, fig. 19)
narrow, whitish, the prearcular and costal regions light yellow;
wing apex narrowly darkened ; stigma small, dark brown ; vague,
scarcely evident dark seams along cord, the veins of the radial
field, vein Cux, and a spot between the anal veins at point
of divergence; veins dark brown, yellow in the flavous areas.
Venation: R2 about one-half its length before the fork of R3 + 4;
inner ends of cells R5 and M3 nearly in alignment ; m-cu shortly
before the fork of M ; apical fusion of Cut and 1st A punctif orm.
Abdominal tergites black, the sternites abruptly orange yel-
low ; a conspicuous subterminal black ring ; female genitalia yel-
low horn color; male hypopygium chiefly darkened.
46, 3 Alexander: Philippine Tipulidze, XII 473
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,500 feet, September 5, 1930 (C. F. Clagg) ; holotype, male;
allotype, female ; paratype, female.
The paratype is much smaller than the other types, as shown
by the measurements. By my key to the Philippine species of
Trentepohlia 2 the present species runs to couplet 10, disagreeing
with both included species in the venation and wing pattern.
The fly is most nearly related to T. (M.) luzonensis Edwards and
allied species that have been discussed and keyed under the de-
scription of T. (M.) sequialba sp. no v.
TRENTEPOHLIA (MONGOMA) MAJUSCULA sp. nov. Plate 1, fig. 20.
Male. — Length, about 15 to 16 millimeters; wing, 10 to 10.5.
Female. — Length, about 16 to 16.5 millimeters; wing, 11.3
to 11.5.
Closely allied to T. (M.) xquinigra sp. nov., differing espe-
cially in the larger size and details of coloration.
Mesonotal praeseutum rich fulvous orange, most intense me-
dially, in cases entirely clear, in other specimens (including the
holotype) narrowly blackened on either side at the suture; scu-
tum with an irregular brown area on either lobe; scutellum
chiefly testaceous yellow ; postnotal mediotergite with the central
portion yellow, the posterior margins darkened, the lateral por-
tions again brightened. Pleura yellow to orange yellow. Hal-
teres yellow, the knobs dark brown. Legs long and powerful;
coxae and trochanters concolorous with the pleura ; femora chiefly
light brown, the bases narrowly more yellowish, the tips narrow-
ly blackened, the amount of the latter subequal on all legs ; tibiae
brown, the tips broadly blackened; basitarsi black, the outer
segments paling to brown; femora with scattered black setae
scattered over the entire length; a group of slightly longer and
more erect setae at base of posterior tibiae. Wings (Plate 1,
fig. 20) narrow, whitish, the prearcular and costal regions yel-
low; stigma small, dark brown; wing tip very narrowly in-
fumed ; Cu, the cord and veins of the radial field narrowly and
vaguely seamed with darker ; the usual small dark spot between
anal veins present; veins black, C, Sc, and R more yellowish.
Venation: Veins R3 and R4 strongly divergent; inner end of
cell M3 lying slightly proximad of cell R5, the basal section of
M3 being angulated ; m-cu at or close to the fork of M ; apical
fusion of CUi and 1st A punctiform.
2 Philip. Journ. Sci. 43 (1930) 297-298.
474 The Philippine Journal of Science i»3i
Abdominal tergites chiefly yellow, with a narrow, more or
less broken, black longitudinal stripe; sternites uniformly yel-
low; subterminal segments and male hypopygium black. Ovi-
positor with the bases and valves yellowish horn color.
Mindanao, Davao district, Mount Apo, Mainit River, altitude
6,500 feet; Seliban River, 7,000 feet; Galog River trail, 5,000
to 6,000 feet, September 10 to 12, 1930 (C. F. Clagg) ; holotype,
male; allotype, female; paratypes, 1 male, 1 female.
The relationships are shown by the key to the Philippine
species of Trentepohlia allied to luzonensls, as given under the
definition of T. (M.) xquialba sp. nov.
TRENTEPOHLIA (TRENTEPOHLIA) LiETIPENNIS sp. nov. Plate 1, fi*. 21.
Rostrum and palpi black; antennae with the basal segment
of scape black, the flagellum pale; mesonotal prsescutum and
scutum obscure yellow, unmarked; posterior sclerites of meso-
notum brown; pleura blackened, with a more or less distinct
longitudinal pale stripe on dorsal sternopleurite ; halteres black,
the extreme base of the stem yellow; legs yellow; wings whit-
ish, with a heavy dark brown pattern arranged as in the ornar-
tipennis group; vein R3 straight to slightly concave, the cell
pointed at base; basal abdominal segments reddish yellow, the
remainder blackened.
Male. — Length, about 4.5 millimeters; wing, 5.5.
Female. — Length, about 5.5 to 6 millimeters ; wing, 5 to 5.5.
Rostrum and palpi black. Antennae with the basal segment
of scape black, the pedicel and flagellum pale brownish yellow,
more darkened outwardly; antennae (male) relatively elongate,
if bent backward extending almost to the wing root; flagellar
segments long-cylindrical, the verticils shorter than the seg-
ments. Head brownish gray.
Mesonotal prsescutum and scutum obscure yellow, the scutel-
lum and postnotal mediotergite more infuscated. Pleura dark
brown, with a more or less distinct paler longitudinal stripe on
the dorsal sternopleurite. Halteres black, the extreme base of
the stem yellow. Legs with the coxae blackened; trochanters
obscure yellow; remainder of legs yellow. Wings (Plate 1, fig.
21) whitish, with a heavy dark brown pattern, arranged on the
plan of ornatipennis and allies; very heavy brown areas at the
wing base ; at mid-length of wing, sending extensions to vein M
at origin of Rs and to the fork of Rs along the anterior cord;
cells beyond the cord chiefly darkened, variegated by three white
marginal areas in ends of cells R2, R3, and R4 + R5 ; cubital and
46' 3 Alexander: Philippine Tiptdidse, XII 475
anal cells chiefly clear ; veins Cu and m-cu seamed with brown ;
outer portion of cell 1st A extensively clouded with gray; veins
pale, dark in the inf uscated areas. Venation : Rs a trifle longer
than R 2+3+4; vein R3 straight or very gently concave, the inner
end of the cell thus pointed; second section of M and R5 +
M1+2 subequal and both about equal to the basal section of M1+2;
apical fusion of Cu2 and 1st A slight.
Abdomen with the basal four segments reddish yellow, the
remainder of the abdomen, including the hypopygium and ovi-
positor, black; in female, the lateral margins of the basal seg-
ments more or less darkened.
Mindanao, Davao district, Mount Apo, Galog River, altitude
6,000 feet, September 16 to 26, 1930 (C. F. Clagg) ; holotype,
male ; allotype, female ; paratypes, 1 male, 2 females.
Trentepohlia (Trentepohlia) hetipennis is closely allied to
species such as T. (T.) ornatipennis Brunetti (southwest India),
T. (T.) festivipennis Edwards (Perak), and T. (T.) venusti-
pennis Edwards (Borneo). It differs in the coloration of the
body and the details of wing pattern and venation, falling
closest to ornatipennis in the wing pattern but differing there-
from in the venation of the radial field and coloration of the body.
In the present species, and very possibly in the other species of
the group, the tip of R1+2 is atrophied.
TRENTEPOHLIA (ANCHIMONGOMA) APOICOLA sp. nov. Plate 1, fig. 22.
Head dark gray ; general coloration of mesonotum dark brown,
the humeral region extensively obscure yellow; pleura yellow,
the ventral sternopleurite inf uscated; tibiae with the central
half to three-fifths blackened.
Male. — Length, about 7 to 8.5 millimeters; wing, 7 to 8.
Rostrum dark, the labial palpi yellow; maxillary palpi black.
Antennae black throughout ; flagellar segments with verticils that
exceed the segments. Head dark gray.
Mesonotal praeseutum medially dark brown to black, more in-
tense in front, the humeral region extensively obscure yellow;
posterior sclerites of mesonotum chiefly darkened, the scutellum
obscure yellow. Pleura obscure yellow, the ventral sternopleu-
rite infuscated. Halteres brownish black, the base of the stem
restrictedly obscure yellow. Legs with the coxae and trochanters
yellow ; femora black, the tips broadly and conspicuously snowy
white, the amount subequal on all the legs ; tibiae black, the cen-
tral portion blackened, most extensively on the posterior legs
where about three-fifths of the segment is included ; tarsi white.
476 The Philippine Journal of Science 1931
Wings (Plate 1, fig. 22) grayish, cells C, Sc, and the apex a trifle
darker; veins dark brown, those of the medial field paler. Ve-
nation : SCi ending opposite the cephalic end of R2, Sc2 opposite
the fork of Rs; cell Cu widely open at margin.
Abdomen dark brown, the basal sternites more yellowish;
hypopygium black.
Mindanao, Davao district, Mount Apo, altitude 6,000 to 8,000
feet, August 30 to September 22, 1930 (C. F. Clagg) ; holotype,
male; paratypes, several males.
Trentepohlia (Anchimongoma) apoicola is very close to T.
(A.) niveipes Edwards (Java), differing only in the details of
coloration of the body and the slightly increased amount of black
on the posterior tibiae.
ILLUSTRATIONS
[Legend : a, ssdeagus ; b, basistyle ; d, dististyle ; g ; gonapophysis ; i, interbasal process ; id,
inner dististyle; od, outer dististyle; p, phallosome ; t, tergite.]
Plate 1
Fig. 1. Limonia (Laosa) manobo sp. nov., wing.
2. LiTrwnia (Limonia) bilan sp. nov., wing.
3. Limonia (Limonia) atroaurata sp. nov., wing.
4. Limonia (Limonia) bagobo sp. nov., wing.
5. Limonia (Limonia) subpacata sp. nov., wing.
6. Limonia (Limonia) subprolixa sp. nov., wing.
7. Helius (Helms) procerus sp. nov., wing.
8. Helius (Helius) apoensis sp. nov., wing.
9. Thaumastoptera (Thaumastoptera) maculivena sp. nov., wing.
10. Adelphomyia apoana sp. nov., wing.
11. Adelphomyia paucisetosa sp. nov., wing.
12. Epiphragma (Polyphragma) fuscofasciata sp. nov., wing.
13. Epiphragma (Polyphragma) nigrotibiata sp. nov., wing.
14. Epiphragma (Polyphragma) apoensis sp. nov., wing.
15. Epiphragma (Polyphragma) caninota sp. nov., wing.
16. Epiphragma (Polyphragma) griseicapilla sp. nov., wing.
17. Epiphragma (Polyphragma) angusticrenula sp. nov., wing.
18. Trentepohlia (Mongoma) sequialba sp. nov., wing.
19. Trentepohlia (Mongoma) sequinigra sp. nov., wing.
20. Trentepohlia (Mongoma) majuscula sp. nov., wing
21. Trentepohlia (Trentepohlia) Isetipennis sj). nov., wing.
22. Trentepohlia (Anchimongoma) apoicola sp. nov., wing.
Plate 2
Fig. 23. Limonia (Limonia) bagobo sp. nov., male hypopygium.
24. Limonia (Limonia) subpacata sp. nov., male hypopygium.
25. Limonia (Limonia) subprolixa sp. nov., male hypopygium.
26. Helius (Helius) procerus sp. nov., male hypopygium.
27. Thaumastoptera (Thaumastoptera) maculivena sp. nov., male hy-
popygium.
28. Adelphomyia paucisetosa sp. nov., male hypopygium.
29. Epiphragma (Polyphragma) latitergata sp. nov., male hypopy-
gium.
30. Epiphragma (Polyphragma) nigrotibiata sp. nov., male hypopy-
gium.
31. Epiphragma (Polyphragma) apoensis sp. nov., male hypopygium.
32. Epiphragma (Polyphragma) hastata sp. nov., male hypopygium.
33. Epiphragma (Polyphragma) caninota sp. nov., male hypopygium.
34. Epiphragma (Polyphragma) griseicapilla sp. nov., male hypopy-
gium.
35. Epiphragma (Polyphragma) angusticrenula sp. nov., male hypo-
pygium.
477
Alexander: Philippine Tipulidte, XII. 1
[Philip. Journ. Sci., 46, No. 3.
PLATE 1.
Alexander: Philippine Tipulid/e, XII.]
[Philip. Journ. Sci., 46, No. 3.
PLATE 2.
SECOND SUPPLEMENT TO THE LIST OF THE LOWER
FUNGI OF THE PHILIPPINE ISLANDS x
A BIBLIOGRAPHIC LIST CHRONOLOGICALLY ARRANGED, AND WITH
LOCALITIES AND HOSTS
By C. F. Baker
Late Dean of the College of Agriculture, Los Banos, Philippine Islands
Edited by F. L. Stevens
Charles Fuller Baker Memorial Professor (1930-1931) of Plant Pathology
UREDINALES
PUCCINIACEJE
HEMILEIA CANTHII Berk, and Br.
On Plectronia. Baker, Philip. Agr. & For. 3 (1914) 160; Philip.
Journ. Sci. 13 (1918) 379.
On Plectronia horrida. Ann. Myc. 26 (1928) 419.
HEMILEIA VASTATRIX Berk, and Br.
On Coffea arabica. Baker, Philip. Agr. & For. 3 (1914) 160; Ann.
Myc. 15 (1917) 175; Philip. Journ. Sci. 13 (1918) 379; Ann. Myc.
26 (1928) 419; Philip. Agr. 17 (1928) 45.
On Coffea spp. Philip. Agr. & For. 6 (1917) 251; Phytopath. 9 (1919)
122; Philip. Agr. 15 (1926) 125; Philip. Agr. Rev. 19 (1926) 252.
1 Contribution from the Experiment Station of the College of Agriculture,
Los Banos, Laguna, Philippine Islands. Published with the approval of the
Director of the Experiment Station.
The editor has chosen to print this article as nearly as, possible as it
was left in manuscript by Dean Baker, with the exception of a consider-
able number of added references, rather than to make changes of which
he might not have approved. The arrangement followed is essentially
that of the two earlier Baker lists. Some of the references give no inter-
nal evidence that the fungi in question occur in the Philippines, but the
fact that they were placed in Dean Baker's manuscript makes it presump-
tive that they do so occur. Dean Baker's work on the manuscript ceased
about 1921.
For the convenience of those who will use this list the editor will issue
later a combined index to the fungi of the Philippine Islands.
Dr. G. O. Ocfemia, of the Department of Plant Pathology of the Col-
lege of Agriculture, at Los Banos, rendered valuable assistance in the
gathering of materials for this manuscript.
479
480 The Philippine Journal of Science 1931
HAMASPORA ACUTISSIMA Syd.
On Rubus moluccanus. (Ann. Myc. 15 (1917) 174; 26 (1928) 418.
PUCCINIA CITRATA Syd.
On Andropogon citratus. Baker, Philip. Agr. & For. 3 (1914) 158;
Philip. Journ. Sci. 13 (1918) 379; Ann. Myc. 21 (1923) 93.
PUCCINIA CONGESTA Berk, and Br.
On Polygonum chinensis. Ann. Myc. 15 (1917) 173.
On Polygonum tomentosum. Ann. Myc. 26 (1928) 416.
PUCCINIA ENGLERIANA P. Henn.
On Tabernaemontana campanulata. Ann. Myc. 15 (1917) 173.
PUCCINIA EREBIA Syd.
On Clerodendron minahassae. Ann. Myc. 15 (1917) 172.
On Clerodendron inermis. Ann. Myc. 26 (1928) 416.
PUCCINIA HETEROSPORA Berk, and Curt.
On Sida javensis. Philip. Journ. Sci. 13 (1918) 379; Ann. Myc. 26
(1928) 416.
PUCCINIA KUEHNII (Kruegr.) Butl. [Uredo kuehnii (Krue*.) Wakk. and Went.]
On Saccharum officinarum. Baker, Philip. Agr. & For. 3 (1914) 164;
Philip. Agr. Rev. 11 (1918) 275; Reinking, Philip. Journ. Sci. 15
(1918) 169; Phytopath. 9 (1919) 135; Philip. Agr. Rev. 14 (1921)
430; Ann. Myc. 26 (1928) 417.
PUCCINIA MERRILLII P. Henn.
On Smttax bracteata. (Ann. Myc. 15 (1917) 173.
On Smilax reticulata. Leafl. Philip. Bot. 9 (1925) 3133.
PUCCINIA PAULLULA Syd.
On Amorphophallus eampanulatus. Ann. Myc. 15 (1917) 173; 26
(1928) 417.
PUCCINIA PHILIPPINENSIS Syd.
On Cyperus compressus. Ann. Myc. 15 (1917) 173.
PUCCINIA PURPUREA Cke.
On Andropogon sorghum (Sorghum vulgare, Holcus sorghum). Sac-
cardo, Nuovo Giorn. Bot. Ital. 23 (1916) 16— Los Bafios (Baker
371,7) ; Baker, Philip. Journ. Sci. 5 (1916) 77; Philip. Agr. & For.
5 (1916) 77; Ann. Myc. 15 (1917) 174; Reinking, Philip. Journ.
Sci. 13 (1918) 165; Phytopath. 9 (1919) 137; Ann. Myc. 26 (1928)
417.
On Andropogon halepensis. Ann. Myc. 21 (1923) 93.
PUCCINIA THWAITESII Berk.
On Justieia gendarussa. Baker, Philip. Agr. & For. 3 (1914) 161;
Ann. Myc. 15 (1917) 173; Philip. Journ. Sci. 13 (1918) 379; Ann.
Myc. 26 (1928) 417.
PUCCINIOSTELE CLARKIANA (BarcL) Diet.
On Astilbe philippinensis. Ann. Myc. 15 (1917) 175.
46,3 Baker: Lower Fungi 481
SPHAEROPHRAGMIUM LUZONICUM Yates.
On Albizzia procera. Philip. Journ. Sci. 13 (1918) 379; Ann. Myc. 20
(1922) 66; 26 (1928) 418.
UROMYCES APPENDICULATUS (Pers.) Lk.
On Vigna spp. Philip. Agr. & For. 4 (1914) 164; 5 (1916) 77; Ann.
Myc. 21 (1923) 93.
On Phaseolws spp. Reinking, Philip. Journ. Sci. 13 (1918) 169;
Philip. Agr. 10 (1922) 349.
On Phaseolus mungo. Phytopath. 9 (1919) 132.
UROMYCES DEERINGIAE Syd.
On Deeringia baccata. Ann. Myc. 26 (1928) 414.
UROMYCES LINEARIS Berk, and Br.
On Panicum. Saocardo, Nuovo Giorn. Bot. Ital. 23 (1916) 16 — Los
Baiios (Baker 3736).
On Panicum repens. Ann. Myc. 15 (1917) 172; 26 (1928) 415.
UROMYCES MUCUNAE Rabh.
On Mucuna deeringiana (Stizolobium deeringvanwrn) . Baker, Philip.
Agr. & For. 3 (1914) 164; Philip. Journ. Sci. 13 (1918) 168.
On Mucuna cochinchinensis (M. nwea, Stizolobium niveum). Phyto-
path. 9 (1919) 132; Ann. Myc. 26 (1928) 415.
UROMYCES SOJAE Syd.
On Glycine max (G. hispida, G. sojae). Baker, Philip. Agr. & For.
3 (1914) 161; Ann. Myc. 15 (1917) 172; Philip. Journ. Sci. 13
(1918) 167; Phytopath. 9 (1919) 126.
UROMYCES WEDELIAE P. Henn.
Saccardo, Syll. Fung. 17 (1895) 245; Hennings, Hedwigia (1904)
150 — Japan; Baccarini, Ann. Bot. 4 (1907) tab. 10, f. 9; Sydow, Mo-
nogr. Ured. 2 (1907) 15; Ann. Myc. 15 (1917) 172.
COLEOSPORIACEJE
COLEOSPORIUM EXACI Syd.
On Exacum chironioides. Ann. Myc. 26 (1928) 425.
COLEOSPORIUM KNOXIAE Syd.
On Knoxia corymbosa. Ann. Myc. 26 (1928) 425.
COLEOSPORIUM MERRILLII P. Henn.
On Orchidaceae. Baker, Philip. Agr. & For. 3 (1914) 163.
SCHROETERIASTER CINGENS Syd.
On Bridelia glabrifolia. Ann Myc. 26 (1928) 423.
MELAMPSORACE^
KUEHNEOLA DESMIUM (Berk, and Br.) Arth.
On Gossypium spp. Baker, Philip. Agr. & For. 3 (1914) 161; Philip.
Journ. Sci. 13 (1918) 167.
2G8774 12
482 The Philippine Journal of Science 1931
KUEHNEOLA FICI (Cast.) BvtI.
On Ficus carica. Baker, Philip. Agr. & For. 3 (1914) 161; Philip.
Journ. Sci. 13 (1918) 167; Phytopath. 9 (1919) 124, 128.
On Morus alba (M. albus). Baker, Philip. Agr. & For. 3 (1914)
162; Ann. Myc. 15 (1917) 175.
KUEHNEOLA FICI (Cast.) Butl. f. MORICOLA P. Henn.
On Morus alba (M. albus). Saccardo, Nuovo Giorn. Bot. Ital. 23
(1916) 16— Los Baiios (Baker 3735); Syll. Fung. 17 (1916) 451
(Uredo moricola) ; Syll. Fung. 9 (1916) 334 (Uredo moH) ; Philip.
Journ. Sci. 13 (1918) 168.
PHAKOSPORA PACHYRHIZI Syd.
On Pachyrrhizus erosus (P. angulatus, Carcara erosa). Philip. Agr.
& For. 4 (1914) 163; Ann. Myc. 15 (1917) 175; Philip. Journ.
Sci. 13 (1918) 168; Phytopath. 9 (1919) 131; Ann. Myc. 26 (1928)
422.
PHAKOSPORA PHYLLANTHI Diet.
On Phyllanthus sp. Ann. Myc. 15 (1917) 175.
On Phyllanthus niruri. Ann. Myc. 26 (1928) 423.
UREDINALES IMPERFECTA
AECIDIUM ALCHORNEAE Sacc.
On Alchornea rugosa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
17 — Mount Maquiling (Baker 3786).
AECIDIUM BANOSENSE Syd.
On Vemonia vida-li. Ann. Myc. 26 (1928) 426.
AECIDIUM BLUMEAE P. Henn.
On Blumea balsamifera. Ann. Myc. 15 (1917) 176; 26 (1928) 426.
AECIDIUM CLERODENDRI P. Henn.
On Clerodendron fragrans. Ann. Myc. 15 (1917) 176.
AECIDIUM ELAEAGNI-LATIFOLIAE Petch.
On Elaeagnus phUippinensis. Ann. Myc. 26 (1928) 426.
AECIDIUM FLAVIDUM Berk, and Br.
On Pavetta indica. Leafl. Philip. Bot. 9 (1925) 3133.
AECIDIUM KAERNBACHII P. Henn.
On Ipomoea pes-caprae. Ann. Myc. 15 (1917) 176.
On Lepistemon flavescens. Philip. Journ. Sci. 13 (1918) 378; Ann.
Myc. 26 (1928) 427.
AECIDIUM LAGUNENSE Syd.
On Gymnema tingentis. Ann. Myc. 26 (1928) 427.
AECIDIUM LUZONIENSE P. Henn.
On Phyllanthus sp. Ann. Myc. 26 (1928) 427.
AECIDIUM NUMMULARE Berk.
On Ceropegia sp. Ann. Myc. 26 (1928) 427.
46,3 Baker: Lower Fungi 483
AECIDIUM PAEDERIAE Diet.
On Paederia foetida (P. tomentosa). Ann. Myc. 15 (1917) 176; 26
(1928) 427.
AECIDIUM RHYTISMOIDEUM Berk, and Br.
On Diospyros discolor. Philip. Journ. Sci. 13 (1918) 379.
AECIDIUM UVARIAE-RUFAE P. Henn.
On Uvaria rufa. Ann. Myc. 15 (1917) 176; 26 (192$) 428.
UREDO ARTHRAXONIS-CILLARIS P. Henn.
On Arthraxonis sp. Ann. Myc. 15 (1917) 177.
On Anthraxonis quartiniant. Ann. Myc. 26 (1928) 428.
UREDO CLAOXYLI Sacc.
On Claoxylum sp. Saccardo, Ann. Myc. 13 (1915) 126 — Mount Ma-
quiling (Baker 2787).
UREDO DAVAOENSIS Syd.
On Cyanotis axillaris. Ann. Myc. 26 (1928) 428.
UREDO DESMIUM (Berk, and Br.) Petch.
On Gossypium sp. Philip. Journ. Sci. 13 (1918) 167; Phytopath. 9
(1919) 126.
UREDO DIOSCOREAE (Berk, and Br.) Petch.
On Dioscorea esculenta. Philip. Journ. Sci. 13 (1918) 167; Phytopath.
9 (1919) 124.
UREDO DIOSCOREAE-ALATAE Rac.
On Dioscorea alata. Baker, Philip. Agr. & For. 3 (1914) 161; Ann.
Myc. 15 (1917) 177.
On Dioscorea esculenta. Philip. Journ. Sci. 13 (1918) 167.
On Dioscorea. Phytopath. 9 (1919) 124.
UREDO ERYTHRINAE P. Henn.
On Erythrina indica. Hennings, Ann. Mus. du Congo 5, II, Fasc. iii
(1908) 224— Congo; Saccardo and Trotter, Syll. Fung. 21 (1912)
790; Ann. Myc. 15 (1917) 177.
UREDO FICI Cast.
On Ficus carica Linn. Philip. Journ. Sci. 13 (1918) 167.
UREDO MANILENSIS Syd.
On Tabernaemontana polygama. Ann. Myc. 15 (1917) 177.
UREDO OCHRACEA Diet.
On Commelina. Dietel, Hedwigia 35 (1897) — Brazil; Saccardo and
Sydow, Syll. Fung. 14 (1899) 403.
UREDO OPERCULINAE Syd.
On Operculina turpethum. Ann. Myc. 15 (1917) 177; 26 (1928)
429.
484 The Philippine Journal of Science 1m
UREDO PREMNAE Koord.
On Premna vestita. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 16 —
Los Banos (Baker 8828),
On Premna cumingiana. Ann. Myc. 15 (1917) 177.
UREDO VIGNAE Bres.
On Vigna spp. Baker, Philip. Agr. & For. 3 (1914) 164 [Uromyces
appendiculatus (Pers.) Lk.] ; Baker, Philip. Agr. & For. 5 (1916)
77; Philip. Journ. Sci. 13 (1918) 170; Phytopath. 9 (1919) 139.
On Phaseolus mungo. Ann. Myc. 15 (1917) 177.
On Glycine hispida (G. max, G. sojae). Ann. Myc. 21 (1923) 94.
USTILAGINALES
USTILAGINACEJE
CINTRACTIA AXICOLA (Berk.) Cornu.
On Fimbristylis diphylla. Ann. Myc. 15 (1917) 178.
USTILAGO ANDROPOGONIS-ACICULATI Petch.
On Andropogon aciculatus. Ann. Myc. 26 (1928) 430.
USTILAGO FLAGELLATA Syd.
On RottboeUia exaltata. Ann. Myc. 15 (1917) 178; 26 (1928) 430.
USTILAGO ISACHNES Syd.
On Isachne miliacea. Ann. Myc. 15 (1917) 178; 26 (1928) 430.
USTILAGO MANILENSIS Syd.
On Panicivm indicum. Ann. Myc. 15 (1917) 178.
USTILAGO SCITAMINEA (Rabh.) Syd. (Ustilago sacdiari Rabh.)
On Saccharum officinarum. Philip. Agr. Rev. 1 (1908) 295; 2 (1909)
14; Baker, Philip. Agr. & For. 3 (1914) 164; 5 (1916) 76; Philip.
Journ. Sci. 13 (1918) 169; Philip. Agr. Rev. 11 (1918) 275; Phy-
topath. 9 (1919) 135; Philip. Agr. Rev. 14 (1921) 428; 18 (1925)
562.
USTILAGO SORGHI (Lk.) Pass.
On Andropogon sorghum (Sorghum vulgaret Holcus sorghum). Philip.
Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919) 137.
USTILAGO TONGLINENSIS Tracy and Earle.
On Ischaemum aristatum. Ann. Myd. 15 (1917) 178; 26 (1928) 430.
TILLETIACE^
ENTTLOMA ORYZAE Syd.
On Oryza sativa. Baker, Philip. Agr. & For. 4 (1914) 163; Philip.
Journ. Sci. 13 (1918) 378; Phytopath. 9 (1919) 131.
PERISPORIALES
ERYSIPHACE.E
PHYLLACTINIA SUFFULTA (Rebent.) Sacc.
On Morns alba. Philip. Agr. & For. 4 (1914) 162; Philip. Journ. Sci.
13 (1918) 168; Phytopath. 9 (1919) 128.
46, 3 Baker: Lower Fungi 485
PERISPORIACEiE
ACTINODOTHIS PIPERIS Syd.
On Piper retrofractwn. Baker, Leafl. Philip. Bot. 7 (1914) 2451;
Theissen and Sydow, Ann. Myc. 13 (1915) 255; Philip. Journ. Sci.
12 (1917) 374; Ann. Myc. 15 (1917) 223; 26 (1928) 439.
BALLADYNA VELUTINA (Berk, and Curt.) y. Hoehnel.
On Plectronia didyma. Ann. Myc. 15 (1917) 180.
DIMERINA GRAFFII Syd.
On Meliola micromera Syd. on Gmelia philippinensis. Ann. Myc. 15
(1917) 199.
DIMERIUM TAYABENSE Yates.
On Momordica sp. Ann. Myc. 20 (1928) 69; Philip. Journ. Sci. 12
(1917) 362.
MELIOLA AFFINIS Syd.
On Memecylon sp. Philip. Journ. Sci. 12 (1917) 362.
MELIOLA ALIENA Syd.
On fallen branches. Ann. Myc. 15 (1917) 181.
MELIOLA ALSTONIAE Koord.
On Alstonia scholaris. Ann. Myc. 15 (1917) 181.
On Alstonia. Leafl. Philip. Bot. 9 (1925) 3133.
MELIOLA ARACHNOIDEA Speg.
On Triumfetta sp. Ann. Myc. 15 (1917) 182.
MELIOLA ARUNDINIS Pat.
On Phragmites vulgaris. Ann. Myc. 15 (1917) 182; Leafl. Philip. Bot.
9 (1925) 3133.
On Saccharum officinarum. Philip. Agr. & For. 5 (1916) 343; Philip.
Agr. Rev. 11 (1918) 275; Philip. Journ. Sci. 13 (1918) 169; Phy-
topath. 9 (1919) 136; Philip. Agr. Rev. 14 (1921) 431.
MELIOLA BAKERI Syd.
On Tetrastigma sp. Ann. Myc. 14 (1916) 355; 15 (1917) 182; Leafl.
Philip. Bot. 9 (1925) 3134.
MELIOLA CALLICARPAE Syd.
On Callicarpa cana. Ann. Myc. 15 (1917) 182.
On CaMicarpa sp. Philip. Journ. Sci. 13 (1918) 362.
MELIOLA CALLISTA Rehm.
On Premna odorata. Ann. Myc. 15 (1917) 183.
MELIOLA CITRICOLA Syd.
On Citrus sp. Ann. Myc. 15 (1917) 183; Reinking, Philip. Agr. 9
(1920) 138; Ann. Myc. 21 (1923) 96.
MELIOLA CLERODENDRICOLA P. Hemw
On Clerodendron sp. Philip. Journ. Sci. 13 (1918) 363.
486 The Philippine Journal of Science 1931
MELIOLA COOKEANA Spegr. var. SACCARDOI Syd.
On Litsea mollis. Sydow, Ann. Myc. 170— Chile (1904); Saccardo,
Syll. Fung. 17 (1905) 546.
On Litsea glutinosa. Ann. Myc. 15 (1917) 184.
MELIOLA CYLINDROPHORA Rehm.
On Guioa perrottetii. Ann. Myc. 15 (1917) 184.
On I tea macrophylla. Ann. Myc. 21 (1923) 95.
MELIOLA DESMODII Karst. and Roum.
On Desmodiwm pulchellum. Ann. Myc. 15 (1917) 185.
MELIOLA DICHOTOMA Berk, and Cke.
On Phragmitis karka. Ann. Myc. 15 (1917) 185.
MELIOLA ELMERI Syd.
On Pittosporum pentandrum. Ann. Myc. 15 (1917) 185.
On Pittosporum sp. Ann. Myc. 21 (1923) 96.
MELIOLA GYMNOSPORIAE Syd.
On Gymnospora spinosa. Philip. Journ. Sci. 13 (1918) 363.
MELIOLA HEWITTIAE Rehm.
On Hewittia sublobata. Philip. Journ. Sci. 12 (1917) 362; Ann. Myc.
15 (1917) 186; 21 (1923) 96.
MELIOLA HYPTIDIS Syd.
On Hyptis swaveolens. Ann. Myc. 15 (1917) 186; Leafl. Philip. Bot. 9
(1925) 3134.
MELIOLA INTRICATA Syd.
On Scirpus grossus. Ann. Myc. 15 (1917) 186.
MELIOLA MACARANGAE Syd. (Meliola apayaoensis Yates.)
On Macaranga tanarius. Ann. Myc. 15 (1917) 188; Philip. Journ.
Sci. 13 (1918) 364; Ann. Myc. 20 (1922) 67.
MELIOLA MANGIFERAE Earle.
On Mangifera mdica. Baker, Philip. Agr. & For. 4 (1914) 162; Ann.
Myc. 15 (1917) 189; Philip. Journ. Sci. 13 (1918) 363; Phytopath.
9 (1919) 127; Ann. Myc. 21 (1923) 97.
MELIOLA MERREMIAE Rehm.
On Merremia hederacea. Ann. Myc. 15 (1917) 190.
MELIOLA MERRILLII Syd.
On Cissus sp. Ann. Myc. 15 (1917) 190.
MELIOLA MITRAGYNES Syd.
On Mitragyne rotundifolia. Philip. Journ. Sci. 13 (1918) 363.
MELIOLA PANICI Earle.
On Rottboellia exaltata. Ann. Myc. 26 (1928) 431.
46, 3 Baker: Lower Fungi 487
MELIOLA PARENCHYMATICA Gaill.
On Sapindus sp. Ann. Myc. 15 (1917) 191.
MELIOLA PERPUSILLA Syd.
On Tylophora perrottetii. Ann. Myc. 15 (1917) 191.
On Tylophora floribunda. Leafl. Philip. Bot. 9 (1925) 3134.
MELIOLA PIPERINA Syd.
On Pi^r sp. Ann. Myc. 14 (1916) 358; 15 (1917) 191; Leafl. Philip.
Bot. 9 (1925) 3134.
MELIOLA POLYTRICHA Kalch. and Cke.
On Ardisia. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 17 — Los
Banos (Baker 3832).
MELIOLA QUADRISPINA Rac.
On Merremia umbellata. Ann. Myc. 15 (1917) 191.
MELIOLA SANDORICI Rehm.
On Sandoricum koetjape. Ann. Myc. 15 (1917) 192; Leafl. Philip.
Bot. 9 (1925) 3134.
MELIOLA SIDAE Rehm.
On Sida carpinifolia. Ann. Myc. 15 (1917) 192.
On Sida acuta. Leafl. Philip. Bot. 9 (1925) 3134.
MELIOLA SUBSTENOSPORA v. Hoehn. f. ROTTBOELLIAE Rehm.
On Rottboellia exaltata. Leafl. Philip. Bot. 9 (1925) 3134.
MELIOLA TAMARINDI Syd.
On Tamarindus indica. Ann. Myc. 15 (1917) 192; Philip. Journ. Sci.
13 (1918) 363.
MELIOLA TELOSMAE Rehm.
On Telosma sp. Ann. Myc. 15 (1917) 192.
MELIOLA UNCARIAE Rehm.
On Uncaria perrottetii. Ann. Myc. 15 (1917) 193.
PARODIELLA GRAMMODES (Kze.) Cooke.
Australian Fungi (1892) 301.
On Desnwdium triflorum. Philip. Journ. Sci. 13 (1918) 371.
CAPNODIACEiE
AITHALODERMA CLAVATISPORUM Syd.
On Psidium guajava. Baker, Philip. Agr. & For. 4 (1914) 163; Phy-
topath. 9 (1919) 133.
On Antidesma buniup. Philip. Journ. Sci. 12 (1917) 373.
On Ixora sp. Ann. Myc. 15 (1917) 179.
On Chrysophyllum oliviformis. Ann. Myc. 21 (1923) 97.
CAPNODIUM FOOTII Berk, and Desm.
On Cocos nucifera. Baker, Philip. Agr. & For. 4 (1914) 160; Philip.
Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 121.
488 The Philippine Journal of Science 1&31
FUMAGO VAGANS Pers.
On Andropogon sorghum (Sorghum vulgar 'e, Holcus sorghum). Ann.
Myc. 15 (1917) 264; Philip. Journ. Sci. 13 (1918) 165; Phytopath.
9 (1919) 138.
LIMACINIA BISEPTATA Sacc.
On Macaranga sp. Saccardo, Ann. Myc. 13 (1915) 127— Los Banos
(Baker 2583).
LIMACINULA MALLOTI Rehm.
On Mallotus philippinensis. Philip. Agr. & For. 4 (1914) 164.
MICROXYPHIUM DUBIUM Sacc.
On Pinanga. Saccardo, Ann. Myc. 13 (1915) 127— Los Banos (Re-
yes, comm. Baker 81).
HEMISPHAERIALES
MICROTHYRIACEiE
ASTERINA BREYNIAE Syd. (Asterina breyniae Yates.)
On Breynia cernua. Ann. Myc. 15 (1917) 242; Philip. Journ. Sci. 12
(1917) 370; Ann. Myc. 20 (1922) 71.
ASTERINA CAPPARIDIS Syd. and Butl.
On Capparis micracantha. Philip. Journ. Sci, 12 (1917) 370; Ann.
Myc. 26 (1928) 439.
On Capparis horrida. Ann. Myc. 15 (1917) 243.
On Capparis irosinensis. Leafl. Philip. Bot. 9 (1925) 3137.
ASTERINA CASSIAE Syd.
On Phyllanthus reticulatus. Ann. Myc. 15 (1917) 245.
On Cuestis diffusa. Ann. Myc. 21 (1923) 103.
ASTERINA COLLICULOSA Speg.
On Eugenia jambolana. Philip. Journ. Sci. 12 (1917) 370.
ASTERINA DECIPIENS Syd.
On Champereia manillana. Ann. Myc. 15 (1917) 245; Philip. Journ.
Sci. 13 (1918) 372.
ASTERINA DILLENIAE Syd.
On Dillenia philippinensis. Ann. Myc. 15 (1917) 244.
ASTERINA ELMERI Syd.
On Champereia manillana. Philip. Journ. Sci. 12 (1917) 370; Ann.
Myc. 21 (1923) 103.
On Champereia cumingiana. Ann. Myc. 15 (1917) 245.
On Champereia sp. Leafl. Philip. Bot. 9 (1925) 3137.
ASTERINA GMELINAE Sacc.
On Gmelina. Saccardo, Nuovo Giorn. Bot. Ital. 23, (1916) 17 Los
Banos (Baker 3763).
46> 3 Baker: Lower Fungi 489
ASTERINA LAWSONIAE P. Henn. and Nym.
On Lawsonia inermis. Baker, Philip, Agr. & For. 4 (1914) 162;
Ann. Myc. 15 (1917) 244.
ASTERINA LAXIUSCULA Syd.
On Sideroxylon sp. Ann. Myc. 15 (1917) 244.
On Sideroxylon ferrunginewm* Philip. Journ. Sci. 13 (1918) 372.
ASTERINA LOBATA Syd.
On unknown host. Ann. Myc. 15 (1917) 244.
ASTERINA OPPOSITA Syd.
On Heynea sumatrana. Ann. Myc. 15 (1917) 245.
ASTERINA PIPTURI Syd.
On Pipturus arborescens. Ann. Myc. 14 (1916) 366; 15 (1917) 245;
Leafl. Philip. Bot. 9 (1925) 3137.
ASTERINA PUSILLA Syd.
On Premna sp. Ann. Myc. 15 (1917) 244.
ASTERINA SPONIAE Rac.
On Trema orientalis. Philip. Journ. Sci. 12 (1917) 370.
On Trema amboinensis. Ann. Myc. 15 (1917) 244.
On Trema sp. Ann. Myc. 21 (192*3) 103; Leafl. Philip. Bot. 9 (1925)
3137.
MORENOELLA MEMECYLI Syd.
On Memecylon lanceolatumi. Ann. Myc. 15 (1917) 251; Philip. Journ.
Sci. 12 (1917) 372.
On Memecylon subfurfuraceum. Ann. Myc. 21 (1923) 104.
TRICHOTHYRIUM ORBICULARE Syd.
On Meliola sp. Ann. Myc. 15 (1917) 236.
On Ficus ulmifolia. Leafl. Philip. Bot. 9 (1925) 3136.
SEYNESIA ALSTONIAE Rehm.
On Alstonia macrophylla. Ann. Myc. 16 (1918) 221.
SEYNESIA IPOMOEAE Syd.
On Merremia sp. Ann. Myc. 15 (1917) 239.
ASTERINELLA CALAMI Syd.
On Calamus sp. Ann. Myc. 15 (1917) 248; Philip. Journ. Sci. 13
(1918) 375.
ASTERINELLA LUZONENSIS Syd.
On Shorea sp. Philip. Journ. Sci. 13 (1918) 376.
ASTERINELLA OBESA Syd.
On Canarium sp. Ann. Myc. 15 (1917) 247; 21 (1923) 104.
ASTERINELLA STUHLMANNI (Henn.) Theiss.
On Ananas comosus (A. sativus, A. sativas, Ananassa sativa). Baker,
Philip. Agr. & For. 5 (1916) 73— Los Bafios; Ann. Myc. 15 (1917)
247; Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919) 115;
Ann. Myc. 21 (1923) 104.
490 The Philippine Journal of Science 1931
LEMBOSIA CONGREGATA Syd.
On Rhododendron schadenbergii. Rehm, Leafl. Philip. Bot. 8 (1915)
2931 — Mount Banahao (A. S. Cruz, comm. Baker 2981).
LEMBOSIA CRUSTACEA (Cke.) Theiss.
Cooke, Grevilea 14 (1915) 13 (Asterina) ; Saccardo, Syll. Fung.
9 (1891) 380 (Asterina); Theissen, Ann. Myc 11 (1891) 432;
Baker, Leafl. Philip. Bot. 6 (1914) 2137 (Morenoella breviuscula) .
On Rhododendron schadenbergii. Rhem, Leafl. Philip. Bot. 8 (1915)
2931 — Mount Banahao (Catalan, comm. Baker 2921).
On Rhododendron sp. Ann. Myc. 15 (1917) 249.
LEMBOSIA EUGENIAE Rehm.
On Eugenia. Rehm, Leafl. Philip. Bot. 8 (1915) 2932— Hills back of
Paete, Luzon (Baker 3137a).
On Eugenia calubcub. Ann. Myc. 15 (1917) 249.
LEMBOSIA JAVANICA (Pat.) Rac.
On Nipa fruticans. Philip. Agr. & For. 4 (1914) 163.
LEMBOSIA PANDANI (Rostr.) Theiss.
On Pandanus. Rostrup (Asterina pandani) ; Theissen, Ann. Myc.
(1913) 457; Syll. Fung. 17 (1913) 881; Rehm, Leafl. Philip. Bot.
8 (1915) 3932— Hills back of Paete (Baker 3113b).
On Pandanus copelandi. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
23— Hills back of Paete (Baker 3789).
LEMBOSIA POTHOIDEI Rehm.
On Pothoideum lobbianum. Leafl. Philip. Bot. 9 (1925) 3137.
MERRILLIOPELTIS CALAMI P. Henn.
On Calamus. Rehm, Leafl. Philip. Bot. 8 (1916) 2945 — Mount Maqui-
ling (Baker 2739, 3189); Philip. Journ. Sci. 12 (1917) 377.
MERRILLIOPELTIS DAEMONOROPSIS Syd.
On Daemonorops. Rehm, Leafl. Philip. Bot. 8 (1916) 2945 — Mount
Maquiling (Reyes, comm. Baker 3343).
MERRILLIOPELTIS HOEHNELII Rehm.
On Dinochloa and Arenga saccharifera. Rehm, Leafl. Philip. Bot. 8
(1916) 2945 — Mount Maquiling (Baker 2189) ; Los Banos (Reyes,
comm. Baker 3371).
TRICHOTHYRIACEjE
Theissen, Beih. Bot. Centralbl. 32 (1914) 14.
GILLETIELLA LATEMACULANS Rehm.
On Arenga saccharifera. Philip. Agr. & For. 4 (1914) 158.
LORANTHOMYCES SORDIDULA (Lev.) v. Hoehn.
On Loranthus haenkeani. Baker, Leafl. Philip. Bot. 6 (1914) 2115;
7 (1914) 2468; Ann. Myc. 15 (1917) 236.
On Loranthus sp. Ann. Myc. 21 (1923) 99.
46, 3 Baker: Lower Fungi 491
HEMISPHAERIACEiE
MICROPELTIS AERUGINASCENS Rehm.
On Rourea erecta. Ann. Myc. 15 (1917) 231.
DICTYOTHYRIELLA MUCOSA Syd. (Micropenis mucosa Syd.)
On Coffea excelpa. Baker, Philip. Agr. & For. 5 (1916) 75 — Los Ba-
nos; Ann. Myc. 14 (1916) 364; Philip. Journ. Sci. 13 (1918) 167;
Phytopath. 9 (1919) 122.
MICROTHYRIELLA PHILIPPINENSIS Syd.
On Lepisanthes schizolepis, Evonymus japonicus, Bauhinia cumingiana.
Ann. Myc. 15 (1917) 235.
MICROTHYRIELLA LATEMACULANS (Rehm) Theiss and Syd.
Baker, Leafl. Philip. Bot. 7 (1914) 2443 (Gillettiella) ; Baker, Philip.
Agr. & For. 3 (1914) 158; Theissen and Sydow, Ann. Myc. 8
(1915) 254.
MYIOCOPRELLA BAKERI Sacc.
On Aspidium. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 17 — Paete,
Laguna Province {Baker 3829).
MYIOCOPRON BAKERIANUM Rehm.
On Passiflora quadrangularis. Baker, Philip. Agr. & For. 4 (1914)
163.
MICROPELTELLA CONSIMILIS Rehm.
On Cryptocarya sp. Ann. Myc. 15 (1917) 229.
HYPOCREALES
HYPOCREACE^S
BROOMELLA ZEAE Rehm.
On Zea mays. Rehm, Leafl. Philip. Bot. 8 (1915) 2923 — Los Bafios
(Raimundo, comm. Baker 19 9 U) ; Baker, Philip. Agr. & For. 5
(1916) 78; Philip. Journ. Sci. 13 (1918) 170; Phytopath. 9 (1919)
140.
MEGALONECTRIA PSEUDOTRICHIA (Schw.) Spe*.
On dead bark. Ann. Myc. 15 (1917) 215.
On Hevea brafUiensis. Philip. Journ. Sci. 13 (1918) 167.
NECTRIACEJE
CALONECTRIA COPELANDII P. Henn.
On Orchidaceae. Baker, Philip. Agr. & For. 4 (1914) 163.
CALONECTRIA SULCATA Starb.
Starbaeck, Bih. K. Svensk. Vet. Ak. Handl. 25 (1899) 29; Zimmer-
MANN, Centralbl. Bakter. 7 (1901) 106 (C. meliae).
492 The Philippine Journal of Science 1931
CALONECTRIA HIBISCOLA P. Henn. (Calonectria meliae A. Zimm.)
Saccardo and Sydow, Syll. Fung. 16 (1902) 593; Saccardo, Syll.
Fung. 17 (1905) 810 (C. meliae); Weese, Myc. Centralbl. Apr.-May
(1914).
On Ficm pseudopalma. Rehm, Leafl. Philip. Bot. 8 (1915) 2923 — Los
Bafios (Raimundo, comm. Baker 1397b).
GIBBERELLA SAUBINETII (Mont.) Sacc.
On Hibiscus esculentus. Baker, Philip. Agr. & For. 4 (1914) 161.
On Panicum sp. Leafl. Philip. Bot. 9 (1925) 3135.
LISEA REVOCANS Sacc.
On Imperata cylindrica. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
23— Los Bafios (Baker 3738); Ann. Myc. 15 (1917) 214.
NECTRIA BAINII Massee.
On Theobroma cacao. Philip. Agr. & For. 4 (1915) 164; Phytopath.
9 (1919) 138.
NECTRIA BAINII Massee var. HYPOLEUCA Sacc.
On Theobroma cacao. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
23-— Los Banos (Baker 3887) ; Baker, Philip. Agr. & For. 5 (1916)
77; Philip. Journ. Sci. 13 (1918) 169.
NECTRIA DISCOPHORA Mont.
Montagne, Syll. gen. sp. Crypt, n. 782 (Sphaeria) (1856) ; Saccardo,
Syll. Fung. 2 (1883) 488; Zimmermann, Centralbl. Bakter. 7 (1901)
106 (N. striatospora) ; Saccardo and Sydow, Syll. Fung. 16 (1902)
1140 (N. striatospora); Weese, Gaehrungsphys. 6 (1902) 114r-121;
De Jonge, Rec. Trav. Botan. Neerl. 6 (1909) tab. 3, f. 14-17 (N.
striatospora) .
NECTRIA SUBFURFURACEA P. Henn. and Nym.
Hennings and Nymann, Monsunia 1 (1899) 162.
On dead fallen branches. Rehm, Leafl. Philip. Bot. 8 (1915) 2922—
Mount Maquiling (Baker 2132).
NECTRIA TJIBODENSIS Penz. and Sacc. var. GLIRICIDIAE Rehm.
On Gliricidia septum. Rehm, Leafl. Philip. Bot. 8 (1915) 2922 — Los
Bafios (Raimundo, comm. Baker 1496).
OPHIONECTRIA ERINACEA Rehm.
On Bambusa blumeana. Baker, Philip. Agr. & For. 4 (1914) 158.
OPHIONECTRIA THEOBROMAE (Pat.) Duss.
On Theobroma cacao. Baker, Philip. Agr. & For. 4 (1914) 164;
4 (1915) 165; 5 (1916) 77; Philip. Journ. Sci. 13 (1918) 169;
Phytopath. 9 (1919) 139.
PARANECTRIA LUXURIANS Rehm.
On Meliola maesae and Panicum. Rehm, Leafl. Philip. Bot. 8 (1915)
2924 — Los Banos (Baker 699b) ; (Eladio Sdblan, comm. Baker
2882b) ; (Baker 2800).
4«» 3 Baker: Lower Fungi 493
TRICHONECTRIA BAMBUSICOLA Rehm.
On Bambusa. Baker, Philip. Agr. & For. 3 (1914) 159.
CLAVICIPTiE
EPICHLOE WARBURGIANA P. Magn.
On Donax cannaeformis. Ann. Myc. 15 (1917) 216.
HYPOCRELLA DISCOIDEA (Berk, and Br.) Sacc.
Berkeley and Broome, Journ. Linn. Soc. Bot. 14 (1873) 113 (Hypo*
crea); Saccardo, Michelia 1 (1873) 322; Saccardo, Syll. Fung. 2
(1883) 580; Massee, Kew Bull. 174 (1899) (H. zingiberis) ; Sac-
cardo and Sydow, Syll. Fung. 16 (1902) 603; Hennings, Hedwigia
(1902) 142 (H. zimmerrnanniana) ; Saccardo, Syll. Fung. 17 (1905)
817 (H. zimmerrnanniana) ; Koorders, Bot. Untersuch. (1907) 179
(H. grewiae) ; Saccardo and Trotter, Syll. Fung. 22 (1913) 503
(H. grewiae) ; Petch, Ann., Roy. Bot. Gard. Peradeniya 5 (1914)
526 (Aschersonia-stage: A. samoensis Henn.).
HYPOCRELLA MOLLII Koord.
Koorders, Bot. Untersuch. (1907) 179; v. Hoehnel, Sitz Kais. Akad.
Wiss. Wien 118 (1909) Abth. 1. p. 311 (H. cretacea) ; Saccardo
and Trotter, Syll. Fung. 22 (1913) 504 (H. mollii) ; 506 (H. cre-
tacea) ; Petch, Ann. Roy. Bot. Gard. Peradeniya 5 (1914) 526 (As-
chersonia-stage ; A. confluens Henn.).
HYPOCRELLA RECIBORSKII A. Zimm.
Zimmermann, Centralbl. f. Bakt. 7 (1901) 875; Hennings, Engler's
Bot. Jahrb. 38 (1905) 13 (H. wameckiana) ; Saccardo, Syll. Fung.
17 (1905) 818; Raciborski, Bull. Akad. Sci. Cracovie (1906) 909
(Barya salaccensis) ; Baker, Leafl. Philip. Bot. 6 (1914) 2100; 7
(1914) 2451 (H. salaccensis) ; Petch, Ann. Roy. Bot. Gard. Pera-
deniya 5 (1914) 527.
HYPOCRELLA REINECKIANA P. Henn.
Hennings, Engler's Bot. Jahrb. 23 (1896) 286; Patouillard, Ann.
Bot. Jard. Buitenzorg Suppl. 1 (1897) 125 (H. pernettyae) ; Raci-
borski, Bull. Acad. Sci. Cracovie (1906) 907 (H. globosa) ; Baker,
Leafl. Philip. Bot. 7 (1914) 2451 (H. pemettiae) ; Petch, Ann. Roy.
Bot. Gard. Peradeniya 5 (1914) 524 (Aschersonia-stage ; A. sclero^
toides Henn.).
HYPOCRELLA SALACCENSIS (Rac.) Petch.
On Premna odorata. Ann. Myc. 15 (1917) 215.
HYPOCRELLA SCHIZOSTACHYH P. Henn.
On Schizostachyum. Philip. Journ. Sci. 13 (1918) 376.
OPHIODOTHIS THANATOSPORA (Ley.) Rac.
Leveille, Ann. Sci. Nat. No. 248 (Dothidea) (1845) ; Raciborski, Bull.
Sci. Ak. Crac. (1906) 904.
On Centotheca latifolia. Rehm, Leafl. Philip. Bot. 8 (1915) 2924 —
Mount Maquiling (Baker 2219).
494 The Philippine Journal of Science 1531
USTILAGINOIDEA OCHRACEA P. Henn.
On Panicum sp. Leafl. Philip. Bot. 9 (1925) 3138.
USTILAGINOIDEA VIRENS (Cke.) Takahashi.
On Oryza saliva. Baker, Philip. Agr. & For. 4 (1914) 163; 5 (1916)
75; Ann. Myc. 15 (1917) 217; Philip. Journ. Sci. 13 (1918) 168,
376; Phytopath. 9 (1919) 130; Philip. Agr. Rev. 19 (1926) 240.
DOTHIDEALES
DOTHIDIACE^E
AUERSWALDIA EXAMINANS (Mont, and Berk.) Sacc.
Montagne and Berkeley, Lond. Journ. Bot. 1 (1842) 156 (Sphaeria) ;
PL Javan. (1842) 520 (Dothidea) ; Cooke, Grevilea 13 (1842) 61;
Philip. Agr. & For. 4 (1914) 158; Baker, Leafl. Philip. Bot. 6
(1914) 2101; Leafl. Philip. Bot. 7 (1914) 2452; Theissen and Sy-
dow, Ann. Myc. 13 (1915) 298.
On Hevea brasiliensis. Ann. Myc. 21 (1923) 102.
AUERSWALDIA GIGANTOCHLOAE Rehm.
Theissen and Sydow, Ann. Myc. 13 (1915) 301.
BALANSIA CLAVICEPS Spe«.
On Centotheca latifolia. Ann. Myc. 15 (1917) 216.
DOTHIDELLA GIGANTOCHLOAE (Rehm) Theiss. and Syd.
On Gigantochloa scribneriana. Rehm, Leafl. Philip. Bot. 6 (1914)
2223 (Scirrhia); Baker, Leafl. Philip. Bot. 7 (1914) 2462 (Scirr-
hia); Theissen and Sydow, Ann. Myc. 13 (1915) 320; 15 (1917)
223.
ELMEROCOCCUM ORBICULA Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2102 (Darwiniella) ; Theissen
and Sydow, Ann. Myc. 13 (1915) 282.
HETERODOTHIS LEPTOTHECA Syd.
Sydow, Philip. Journ. Sci. § C 9 (1914) 170; Baker, Leafl. Philip. Bot.
7 (1914) 2454; Theissen and Sydow, Ann. Myc. 13 (1915) 190 is a
lichen and = Phylloporina phyllogena Muell.-Arg.
PSEUDOTHIS PTEROCARPI Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2102 (Dothidea); Theissen and
Sydow, Ann. Myc. 13 (1915) 339.
POLYSTOMELLAGEiE
AULACOSTROMA PALAWANENSE Syd.
On Pandanus tectorius. Baker, Leafl. Philip. Bot. 7 (1914) 2453;
Theissen and Sydow, Ann. Myc. 13 (1915) 256; 15 (1917) 223.
46* 3 Baker: Lower Fungi 495
ELLISIODOTHIS PANDANI Syd.
On Pandanus luzonensis. Sydow, Ann. Myc. (1914) 565 — Angat, Bu-
lacan Province; Theissen and Sydow, Ann. Myc. 13 (1915) 247.
ELLISIODOTHIS REHMIANA Theiss. and Syd.
On Dioscorea esculenta. Baker, Lean. Philip. Bot. 7 (1914) 2460
(Phyllachora) ; Theissen and Sydow, Ann. Myc. 13 (1915) 248;
Philip. Journ. Sci. 13 (1918) 167.
HYSTEROSTOMELLA LETRACERAE (Rud.) v. Hoehnel.
Rud., Linnaea 4 (1829) 118 (Phacidium) ; 5 (1830) 551 (Phacidium) ;
Saccardo, Syll. Fung. 8 (1889) 748 (Coccomyces) ; Ellis and Ever-
HART, Journ. Myc. 10 (1904) 167 (Harknessia) ; Baker, Leafl. Philip.
Bot. 7 (1914) 2497; Theissen and Sydow, Ann. Myc. 13 (1915)
224.
HYSTEROSTOMELLA SPURCARIA (Berk, and Br.) v. Hoehn.
Berkeley and Broome, Fung. Ceyl. (1870) No. 1131 (Rhytisma spur-
carium) ; No. 1132 (Rhytisma constellatum) ; Berkeley and Curtis,
Journ. Linn. Soc. Bot. 14 (1873) 131 (Rhytisma); Saccardo, Syll.
Fung. 8 (1899) 737 (Marchalia) ; v. Hoehnel, Fragm. Myc. 9 (1899)
56.
On Artocarpus communis. Rehm, Leafl. Philip. Bot. 8 (1915) 2932 —
Los Bafios (Baker 2393) ; (Reyes, comm. Baker 2557).
HYSTEROSTOMELLA TETRACERAE (Rud.) v. Hoehn.
On Tetracera sp. Ann. Myc. 15 (1917) 220.
INOCYCLUS PSYCHOTRIAE Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2136; 7 (1914) 2497 (Hysteros-
tomella); Theissen and Sydow, Ann. Myc. 13 (1915) 211.
On Psychotria luzoniensis. Philip. Journ. Sci. 12 (1917) 373; Ann.
Myc. 15 (1917) 220.
MARCHALIA CONSTELLATA (Berk, and Br.) Sacc.
Berkeley and Broome, Journ. Linn. Soc. Bot. 14 (1875) 131 (Rhy-
tisma constellatum and R. spurcarium) ; v. Hoehnel, Fragm. Myc.
9 (1875) No. 448 (HysterostomeUa) ; Saccardo, Syll. Fung. 8 (1899)
737 (Marchalia spurcaria).
On Artocarpus. Theissen and Sydow, Ann. Myc. 13 (1915) 251 —
Philippines, Exsicc: Sydow, Fung. Exot. U03.
On Artocarpus communis. Philip. Journ. Sci. 13 (1918) 165.
MICRODOTHELLA CULMICOLA Syd.
Baker, Leafl. Philip. Bot. 7 (1914) 2454; Theissen and Sydow, Ann.
Myc. 13 (1915) 259.
PALAWANIA COCOES Syd.
On Cocos nucifera. Philip. Agr. & For. 4 (1914) 160; Baker, Leafl.
Philip. Bot. 7 (1914) 2454; Theissen and Sydow, Ann. Myc. 13
(1915) 250; Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919)
122.
496 The Philippine Journal of Science 1931
PALAWANIA GRANDIS (Niessl.) Syd.
Niessl., in Rabh. Fung. Eur. No. 2467 (Microthyrium) ; Winter, Hed-
wigia (1886) 107 (Seynesia); Hennings and E. Nym., (1899) 160
(Seynesia ccdamicola) ; Saccardo and Sydow, Syll. Fung. 16 (1902)
641 (Seynesia calamicola) ; Baker, Leafl. Philip. Bot. 7 (1914)
2454; Theissen and Sydow, Ann. Myc. 13 (1915) 249.
RHIPIDOCARPON JAVANICUM (Pat.) Theiss. and Syd.
Patouillard, Ann. Jard. Buit. Suppl. (1897) 122 (Schneepia) ;
Raciborski, Paras. Alg. und Pilze Javas 2 (1900) 20; (Lembosia);
Saccardo and Sydow, Syll. Fung. 14 (1899) 709 (Parmidaria) ;
Theissen, Ann. Myc. 11 (1912) 453; Baker, Philip. Agr. & For. 3
(1914) 163 (Lembosia); Baker, Leafl. Philip. Bot. 6 (1914) 2138;
7 (1914) 2441 (Lembosia); Theissen and Sydow, Ann. Myc. 13
(1914) 197.
On Nipa fruticans. Rehm, Leafl. Philip. Bot. 8 (1915) 2933— Los Ba-
iios (Reyes, comm. Baker 2548; Mirasol, coram. Baker 1220; Catalan,
coram. Baker 2839); Theissen and Sydow, Ann. Myc. 13 (1915)
197 Exsicc: Rehm, Ascom. 1839 (Java); SydoWi Fung. Exot. 268
(Philippines).
On Psychotria lusoniensis. Ann. Myc. 15 (1917) 220.
SCHNEEPIA HYMENOLEPIDIS (P. Henn.) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2137; 7: 2445 (Parmularia) ; Sy-
dow, Ann. Myc. 13 (1915) 204.
STIGMATODOTHIS PALAWANENSIS Syd.
Baker, Leafl. Philip. Bot. 7 (1914) 2463; Theissen and Sydow, Ann.
Myc. 13 (1915) 264.
ULEOPELTIS BAMBUSINA Syd.
On Bambusa. Sydow, Ann. Myc. 12 (1914) 565— Angat, Bulacan
Province; Theissen and Sydow, Ann. Myc. 13 (1915) 218.
PHYLLACHORACEiE
APIOSPORA APIOSPORA (Dur. and Mtg.) v. Hoehiu
Durand and Montagne, Crypt. Alger, tab. 1 : 492 (Sphaeria) ; Mon-
tagne, Syll. Crypt (1856) No. 809 (Sphaeria); Saccardo, Fung.
Ven. Ser. 2 (1874) 306 (A. montagnei) ; Saccardo, Syll. Fung. 1
(1882) 539 (A. montagnei); Baker, Leafl. Philip. Bot. 6 (1914)
2111 (excl. syn. A. luzonensis) ; Theissen and Sydow, Ann. Myc. 13
(1915) 419 (Apiospora montagnei).
On Bambusa vulgaris. Rehm, Leafl. Philip. Bot. 8 (1916) 2946—Los
Bafios (Reyes, comm. Baker 1895, 1US5).
On Bambusa sp. Ann. Myc. 16 (1918) 223.
APIOSPORA CAMPTOSPORA Pens, and Sacc.
Baker, Leafl. Philip. Bot. 7 (1914) 2463; Theissen and Sydow, Ann.
Myc. 13 (1915) 421; 15 (1917) 225.
On Saccharum officinarum. Philip. Agr. & For. 5 (1916) 343; Philip.
Journ. Sci. 13 (1918) 169; Philip. Agr. Rev. 2 (1918) 276; Phyto-
path. 9 (1919) 136.
46, s Baker: Lower Fungi 497
APIOSPORA CARBONACEA Rehm.
On Schizojstachyum. Rehm, Leafl. Philip. Bot. 8 (1916) 2945 — Mount
Maquiling (Baker 3427a); Leafi. Philip. Bot. 8 (1915) 2945.
APIOSPORA LUZONENSIS P. Henn.
On Bambusa. Baker, Leafi. Philip. Bot. 6 (1914) 2113 (sub Apios-
pora apiospora); Theissen and Sydow, Ann. Myc. 13 (1915) 421;
15 (1917) 225.
CATACAUMA APOENSE (Syd.) Theiss. and Syd.
On Ficus nervosa. Baker, Leafi. Philip. Bot. 6 (1914) 2103 (Phylla-
chora); 7: 2455 (Phyllachora) ; Theissen and Sydow, Ann. Myc. 13
(1915) 2455; 15 (1917) 224.
CATACAUMA ASPIDEUM (Berk.) Theiss. and Syd. f. SPINIFERA (Karst. and Har.)
Theiss. and Syd.
Baker, Leafi. Philip. Bot. 6 (1914) 2106 (Phyllachora fici-minahas-
sae); 2110 (Phyllachora spinif era) ; 7: 2457 (Phyllachora fici-mina-
hassae) ; 2460 (Phyllachora spinif era) ; Theissen and Sydow, Ann.
Myc. 13 (1915) 380.
On Ficus odorata (F. odoratus). Ann. M^c. 15 (1917) 224; Philip.
Journ. Sci. 12 (1917) 374; Ann. Myc. 16 (1918) 215; 21 (1923)
101.
On Ficus validicaudata. Philip. Journ. Sci. 13 (1918) 376.
CATACAUMA ASPIDEUM (Berk.) Theiss. and Syd. f. FICIFULVAE (Koord.) Theiss. and
Syd.
Koorders, Verh. K. Akad. Wet. Amsterdam 2 (1907) 183 No. 4;
Baker, Leafl. Philip. Bot. 6 (1914) 2105 (Phyllachora fici-fulvae) ;
7: 2457 (Phyllachora fici-fulvae); Theissen and Sydow, Ann. Myc.
13 (1915) 381.
On Ficus sp. Philip. Journ. Sci. 12 (1917) 374.
On Ficus odorata (F. odoratus). Ann. Myc. 15 (1917) 224.
CATACAUMA CIRCINATUM (Syd.) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2104 (Phyllachora); 7 (1914)
2455 (Phyllachora); Theissen and' Sydow, Ann. Myc. 13 (1915)
2456.
CATACAUMA ELMERI (Syd.) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2105 (Phyllachora); 7: 2457 (Phyl-
lachora); Theissen and Sydow, Ann. Myc. 13 (1915) 378.
On Ficus sp. Philip. Journ. Sci. 12 (1917) 375.
On Ficus minehassa. Ann. Myc. 15 (1917) 224.
CATACAUMA EURYAE (Rac.) Theiss. and Syd.
Raciborski, Bull. Acad. Cracov. (1909) 377 (Myocopron) ; v. Hoeh-
nel, Fragm. Myc. 7 (1912) No. 305 (Physalospora) ; Baker, Leafl.
Philip. Bot. 6 (1914) 2122 (Physalospora) ; Theissen and Sydow,
Ann. Myc. 13 (1915) 392.
CATACAUMA GARCIAE Theiss. and Syd.
On Ficus garcia. Theissen and Sydow, Ann. Myc. 13 (1915) 381 —
Puerto Princesa, Palawan (Elmer 128^7).
On Ficus sp. Leafl. Philip. Bot. 9 (1925) 3136.
263774 13
498 The Philippine Journal of Science 1931
CATACAUMA INFECTORIUM (Cke.) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2106 (Phyllachora) ; 7 (1914)
2458 (Phyllachora); Theissen and Sydow, Ann. Myc. 13 (1915)
384.
CATACAUMA KAERNBACHII (P. Henn.) Theiss. and Syd.
Hennings, Engl. Bot. Jahrb. 18 Beibl. 44 (1894) 39 (Phyllachora) ;
Syll. Fung. 11: 372 (Phyllachora); Baker, Leafl. Philip. Bot. 6
(1914) 2107 (Phyllachora); 7 (1914) 2458 (Phyllachora); Theissen
and Sydow, Ann. Myc. 13 (1915) 376.
CATACAUMA LAGUNENSE (Syd.) Theiss. and Syd.
On Ficus sp. Baker, Leafl. Philip. Bot. 6 (1914) 2107 (Phylla-
chora); Theissen and Sydow, Ann. Myc. 13 (1915) 378; 15 (1917)
224; Philip. Journ. Sci. 12 (1917) 374.
On Ficus hauili. Ann. Myc. 21 (1923) 101.
CATACAUMA PTEROCARPI (Syd.) Theiss. and Syd.
On Pterocarpus angalensis. Sydow, Ann. Myc. (1912) 40 — South
Africa; Baker, Leafl. Philip. Bot. 6 (1914) 2109 (Phyllachora pte-
rocarpi non Rehm) ; 7 (1914) 2459 (Phyllachora pterocarpi non
Rehm); Theissen and Sydow, Ann. Myc. 13 (1915) 387.
On Pterocarpus indicts, Ann. Myc. 15 (1917) 22'3.
CATACAUMA SANGUINEUM (Rehm) Theiss. and Syd.
On Ficus heterophylla, Baker, Leafl. Philip. Bot. 7 (1914) 2456
(Phyllachora circinata var. sanguinea) ; Theissen and Sydow, Ann.
Myc. 13 (1915) 379; 15 (1917) 224.
CATACAUMA VALSIFORME (Rehm) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2110 (Phyllachora); 7 (1914)
2461 (Phyllachora); Theissen and Sydow, Ann. Myc. 13 (1915)
379.
EXARMIDIUM BLUMEANUM (Rehm) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2110 (Rhopographus) ; 7 (1914)
2462 (Rhopographus); Theissen and Sydow, Ann. Myc. 13 (1915)
425.
MUNKIODOTHIS MELASTOMATA (v. Hoehn.) Theiss. and Syd.
On Melastoma fusca. Baker, Leafl. Philip. Bot. 6 (1914) 2103 (Mun-
kiella) ; 7 (1914) 2454 (Munkiella) ; Theissen and Sydow, Ann.
Myc. 13 (1915) 360; 15 (1917) 223.
PHYLLACHORA AFZELIAE Syd.
On Afzelia bijuga. Philip. Agr. & For. 4 (1914) 163.
PHYLLACHORA CANARI P. Henn.
On Canarium villosum. Philip. Journ. Sci. 12 (1917) 375; Ann. Myc.
16 (1918) 214.
On Canarium sp. Leafl. Philip. Bot. 9 (1925) 3136.
PHYLLACHORA COICIS P. Henn.
On Coix lachryma-jobi. Philip. Journ. Sci. 12 (1917) 375.
46»3 Baker: Lower Fungi 499
PHYLLACHORA CYNODONTIS (Sacc.) Niessl.
On Cynodons dactylis. Ann. Myc. 15 (1917) 227; 26 (1928) 438.
PHYLLACHORA DALBERGIAE Niessl.
On Dalbergia sp. Philip. Journ. Sci. 12 (1917) 375.
PHYLLACHORA DIOSCOREA Schwein.
On Dioscorea sp. Baker, Philip. Agr. & For. 4 (1914) 161; Phyto-
path. 9 (1919) 124.
On Dioscorea esculenta. Philip. Journ. Sci. 13 (1918) 167.
PHYLLACHORA LUZONIENSIS P. Henn.
On Milletia. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 23— Mount
Maquiling (Baker 38tf>) ; Philip. Journ. Sci. 12 (1917) 375; Ann.
Myc. 21 (1923) 102.
On Milletia cavitensis. Ann. Myc. 15 (1917) 225.
PHYLLACHORA MINUTISSIMA (Welw. and Curr.) Sm.
On Pennisetum. Welwitsch and Curr., Trans. Linn. Soc. Bot.
(1868) 285 (Isothea) — Angola; Smith, Journ. Bot. (1898) 179;
Saccardo and Sydow, Syll. Fung. 16 (1902) 623.
PHYLLACHORA ORBICULA Rehm.
On Bambusa blumeana. Philip. Agr. & For. 4 (1914) 158; Ann. Myc.
15 (1917) 227; 16 (1918) 223.
PHYLLACHORA PAHUDIAE Syd.
On Pahudia rhomboidea. Ann. Myc. 15 (1917) 225.
PHYLLACHORA PARKIAE P. Henn.
On Parkia timoriana. Philip. Agr. & For. 4 (1914) 163.
On Parkia javanica. Ann. Myc. 26 (1928) 438.
PHYLLACHORA PHASEOLINA Syd.
Baker, Philip. Agr. & For. 4 (1914) 163.
On Phaseolus sp. Ann. Myc. 15 (1917) 225; Philip. Journ. Sci. 13
(1918) 168.
On Phaseolus calcaratus. Phytopath. 9 (1919) 132.
PHYLLACHORA PONGAMIAE (Berk, and Br.) Petch.
On Pongamia glabra. Philip. Agr. & For. 4 (1914) 163.
On Pongamia pinnata. Philip. Journ. Sci. 12 (1917) 375; Ann. Myc.
21 (1923) 102; 26 (1928) 437.
On Pongamia mitis. Ann. Myc. 15 (1917) 225.
PHYLLACHORA REHMIANA Theiss. and Syd.
On Dioscorea esculenta. Philip. Journ. Sci. 13 (1918) 167.
PHYLLACHORA ROTTBOELLIAE Syd. and Butl.
On Rottboellia exaltata. Leafl. Philip. Bot. 9 (1925) 3136; Ann. Myc.
26 (1928) 438.
PHYLLACHORA ROUREAE Syd.
On Rourea erecta. Ann. Myc. 15 (1917) 226.
500 The Philippine Journal of Science i93i
PHYLLACHORA SACCHARI P. Henn.
On Saccharum officinarum. Philip. Agr. & For. 5 (1916) 343; Philip.
Agr. Rev. 11 (1918) 275; Philip. Journ. Sci. 13 (1918) 169; Phyto-
path. 9 (1919) 134; Philip. Agr. Rev. 14 (1921) 430.
PHYLLACHORA SACCHARI-SPONTANEI Syd.
On Saccharum spontaneum. Ann. Myc. 15 (1917) 226; Philip. Journ.
Sci. 13 (1918) 169; Phytopath. 9 (1919) 134.
PHYLLACHORA SORGHI v. Hoehnel.
On Andropogon halepensis (Sorghum halepensis) var. propinquus.
Ann. Myc. 15 (1917) 226; Philip. Journ. Sci. 13 (1918) 377.
On Andropogon sorghum (Sorghum vulgare, Holcus sorghum, Sorg-
hum sp.). Philip. Journ. Sci. 13 (1918) 165; Ann. Myc. 16 (1918)
214; Phytopath. 9 (1919) 137; Leafl. Philip. Bot. 9 (1925) 3136.
PHYLLACHORA TJANKORREH Rac.
On Dinochloa sp. Ann. Myc. 15 (1917) 228.
On Schizostachyum rotundifoliwn. Ann. Myc. 21 (1923) 102.
PHYLLACHORA YAPENSIS <P. Henn.) Syd.
On Derris elliptica. Philip. Journ. Sci. 12 (1917) 375.
On Derris sp. Ann. Myc. 15 (1917) 225; Leafl. Philip. Bot. 9 (1925)
3136.
RHOPOGRAPHELLA REYESIANA Rchm.
On Schizostachyum sp. Ann. Myc. 15 (1917) 209.
PLACOSTROMA PTEROCARPI (Mass.) Theiss. and Syd.
Massee, Kew Bull. (1912) 257 (Dothidella) ; Baker, Leafl. Philip.
Bot 6 (1914) 2109 (Phyllachora pterocarpi Rehm non Syd.) ; 7
(1914) 2459 (Phyllachora pterocarpi Rehm non Syd.); Theissen
and Sydow, Ann. Myc. 13 (1915) 407.
SCHIZOCHORA ELMERI Syd.
Baker, Leafl. Philip. Bot. 7 (1914) 2462; Theissen and Sydow, Ann.
Myc. 13 (1915) 401.
SCIRRHIA BAMBUSINA Penz. and Sacc.
On Bambusa blumeana.
SCIRRHIA LUZONENSIS P. Henn.
On Bambusa blumeana. Philip. Agr. & For. 4 (1914) 158; Baker,
Leafl. Philip. Bot. 6 (1914) 2111; 7 (1914) 2462; Theissen and Sy-
dow, Ann. Myc. 13 (1915) 418.
SCIRRHODOTHIS BAMBUSINA (Penz. and Sacc.) Theiss. and Syd.
On Schizostachyum acutiflorum. Baker, Leafl. Philip. Bot. 6 (1914)
2111 (Scirrhia); Theissen and Sydow, Ann. Myc. 13 (1915) 416;
Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 23 (Baker 3822).
SCIRRHODOTHIS SERIATA Syd. and Butl.
Baker, Leafl. Philip. Bot. 6 (1914) 2111 (Scirrhia) ; 7: 2463 (Scirrhia) ;
Theissen and Sydow, Ann. Myc. 13 (1915) 416.
46,3 Baker: Lower Fungi 501
APHERODOTHIS ARENGAE (Rac.) Shear.
On Caryota rumphiana var. philippinensis. Ann. Myc. 15 (1917) 228;
Philip. Journ. Sci. 12 (1917) 375; Leafl. Philip. Bot. 9 (1925) 3136.
TRABUTIA ELMERI Theiss. and Syd.
On Ficus banahaensis. Theissen and Sydow, Ann. Mycol. 13 (1915)
353 — Mount Apo, Mindanao (Elmer 10906).
TRABUTIA FICUUM (Niessl.) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2106 (Phyllachora ficuum) ; Theis-
sen and Sydow, Ann. Myc. 13 (1915) 352.
TRABUTIA VERNICOSA Theiss. and Syd.
On Ficus heterophylla. Theissen and Sydow, Ann. Myc. 13 (1915)
353— Mindoro (Merrill 5625).
SPHAERIALES
SORDARIACE^E
SORDARIA ORYZETI Sacc.
On Oryza sativa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 19 —
Los Banos (Baker 3807); Baker, Philip. Agr. & For. 5 (1916) 76;
Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9 (1919) 131.
SPHAERIACEJE
ACANTHOSTIGMA BAMBUSAE v. Hoehn.
V. HOEHNEL, Wiss. Wein. 18: 334.
On Bambusa blumeana. Rehm, Leafl. Philip. Bot. 8 (1916) 3951 —
Los Banos (Baker 2187) ; Mount Maquiling (Baker 3535).
ACERBIA MAYDIS Rehm.
On Zea mays. Rehm, Leafl. Philip. Bot. 8 (1916) 2953 — Los Banos
(Raimundo, comm. Baker 1993) ; Baker, Philip. Agr. & For. 5 (1916)
78; Philip. Journ. Sci. 13 (1918) 170; Phytopath. 9 (1919) 140.
CHAETOSPHAERIA EXIMIA Sacc.
On Cocos nucifera. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 20 —
Los Banos (Baker 3758); Philip. Journ. Sci. 13 (1918) 166; Phy-
topath. 9 (1919) 122.
LASIOSPHAERIA MOLLIS Rehm.
On Bambusa blumeana. Rehm, Leafl. Philip. Bot. 8 (1916) 2952 —
Los Banos (Reyes, comm. Baker 17 3 U).
MELANOMMA MINDORENSE Rehm.
On Arenga saccharifera. Rehm, Leafl. Philip. Bot. 6 (1914) 2202
(Metasphaeria maculans) ; 8 (1916) 2950 — Los Banos (Baker 1876).
MELANOPSAMMA LICHENOIDES Rehm.
On fallen limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2944— Los Banos
(Baker 3067a).
502 The Philippine Journal of Science mi
NEOPECKIA RHODOSTICTA (B. and Br.) Sacc.
On Pandanus. Saccardo, Syll. Fung. 2 (1883) 213 (Herpotrichia) ;
Berlese, Atti Congr. Bot. Genova (1892) 5 (Didymotrichia) ; Rehm,
Leafl. Philip. Bot. 8 (1916) 2946— Los Banos (Reyes, comm. Baker
3U0).
NEOPECKIA RHODOSTICTA (Berk, and Br.) Sacc. var. MAGNIFICA Rehm.
On Pandanus sabutan. Rehm, Leafl. Philip. Bot. 8 (1916) 2947 —
Los Banos (Reyes, comm. Baker 3047).
ROSELLINIA BUNODES (Berk, and Br.) Sacc.
Berkeley and Broome, Fung. Ceylon (1870) No. 1088 (Sphaeria) ;
Saccardo, Syll. Fung. 1 (1882) 254.
On fallen limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2941— Los Banos
(Reyes, comm. Baker); Ann. Myc. 15 (1917) 211.
ROSELLINIA CALAMI P. Henn.
On Bambusa blumeana. Ann. Myc. 15 (1917) 211.
ROSELLINIA COCOES P. Henn.
On Arenga mindorensis. Ann. Myc. 15 (1917) 211.
On Cocos nucifera. Philip. Journ. Sci. 13 (1918) 167; Phytopath. 9
(1919) 122.
ROSELLINIA (TASSIELLA) CRUSTACEA Rehm.
On Schizostachyum.. Rehm, Leafl. Philip. Bot. 8 (1916) 2941 — Los Ba-
nos (Reyes, comm. Baker 3372).
ROSELLINIA DECIPIENS (Rehm) Theiss. and Syd.
Baker, Leafl. Philip. Bot. 6 (1914) 2101 (Auersivaldia) ; Theissen and
Sydow, Ann. Myc. 13 (1915) 300.
ROSELLINIA (TASSIELLA) HORRIDA Rehm.
On dead bark. Rehm, Leafl. Philip. Bot. 8 (1916) 2941— Mount Ma-
quiling (Baker 2909).
ROSELLINIA LAMIPROSTOMA Syd.
On Streblus asper and on dead Daemonorops. Rehm, Leafl. Philip. Bot.
8 (1916) 2942 — Los Banos (Raimundo, comm. Baker 2975) ; Mount
Maquiling (Baker 2720).
ROSELLINIA (CONIMELA) MAQUILINGIANA Rehm.
On fallen limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2942— Mount Ma-
quiling (Reyes, comm. Baker 3347).
ROSELLINIA MEGALOSPERMA Syd.
On Pipturus arborescens. Ann. Myc. 15 (1917) 211.
ROSELLINIA MERRILLII Syd.
On decorticated wood. Ann. Myc. 15 (1917) 211.
ROSELLINIA MOLLERIANA Henn.
Hennings, Hedwigia 13 (1902).
On decorticated wood. Rehm, Leafl. Philip. Bot. 8 (1916) 2942—
Mount Maquiling (Baker 4026).
46,3 Baker: Lower Fungi 503
KOSELLINIA PROCERA Syd.
On Alchomea ru&osa. Rehm, Leafi. Philip. Bot. 8 (1916) 2942— Los
Bafios (Baker 4024).
ROSELLINIA UMBILICATA Sacc.
On bark. Ann. Myc. 15 (1917) 211.
ZIGNOELLA (TREMATOSTOMA) NOBILIS Rehm.
On Citrus nobilis. Rehm, Leafl. Philip. Bot. 8 (1916) 2950 — Los Ba-
nos (Baker 3229); Baker, Philip. Agr. & For. 5 (1916) 74; Phyto-
path. 9 (1919) 119; Reinking, Philip. Agr. 9 (1920-21) 133; Leafl.
Philip. Bot. 8 (1915) 2950; Philip. Journ. Sci. 13 (1918) 166.
CUCURBITARIACE^E
GIBBERA PHILIPPINENSIS Rehm.
On Schizostachyum. Rehm, Leafl. Philip. Bot. 8 (1916) 2946 — Mount
Maquiling (Baker 2896).
NITSCHKEA BAMBUSARUM Rehm.
On Bambusa vulgaris. Rehm, Leafl. Philip. Bot. 8 (1916) 2956 — Los
Banos (Reyes, comm. Baker 1884, 1886).
CORYNELIACEiE
CORYNELIA CLAVATA (L.) Sacc.
On Podocarpus. Rehm, Leafl. Philip. Bot. 8 (1915) 2925 — Mount Ba-
nahao (Copeland, comm. Baker 3639).
On Podocarpus costatus. Ann. Myc. 15 (1917) 178.
TRICHOSPHAERIA BAMBUSICOLA Rehm.
On Bambusa blumeana. Philip. Agr. & For. 4 (1914) 158.
AMPHISPHAERIACEJE
AMPHISPHAERIA ARENGAE Rehm.
On Arenga. Rehm, Leafl. Philip. Bot. 8 (1916) 2947— Los Banos (Re-
yes, comm. Baker 3436).
AMPHISPHAERIA SCHIZOSTACHYI Rehm.
On Schizostachyum. Rehm, Leafl. Philip. Bot. 8 (1916) 2947 — Los Ba-
nos (Baker 1966).
TREMATOSPHAERIA MAQUILINGIANA Rehm.
On Calamus. Rehm, Leafl. Philip. Bot. 8 (1916) 2952— Mount Ma-
quiling (Baker 3420).
TREMATOSPHAERIA MAQUILINGIANA Rehm var. SCHIZOSTACHYI Rehm.
On Schizostachyum. Rehm, Leafl. Philip. Bot. 8 (1916) 2952 — Mount
Maquiling (Baker 3426).
MYCOSPHAERELLACE^E
ASCOSPORA VANILLAE Rehm.
On Vanilla sp. Rehm, Leafl. Philip. Bot. 8 (1916) 2935— Los Banos
(Baker 3079).
504 The Philippine Journal of Science 1931
GUIGNARDIA ARENGAE Rehm.
On Caryota sp. Ann. Myc. 15 (1917) 207.
GUIGNARDIA BAMBUSINA Rehm.
On Bambusa. Rehm, Leafl. Philip. Bot. 8 (1916) 2936 — Los Baiios
(Baker 1898, 1915a).
GUIGNARDIA CREBERRIMA Syd.
On Capparis horrida. Philip. Journ. Sci. 12 (1917) 376; Ann. Myc. 15
(1917) 207; Philip. Journ. Sci. 13 (1918) 377.
GUIGNARDIA DINOCHLOAE Rehm.
On Dinochloa. Rehm, Leafl. Philip. Bot. 8 (1916) 2936— Mount Ma-
quiling (Baker 2189b).
GUIGNARDIA MANIHOTI Sacc.
On Manihot utilissima. Baker, Philip. Agr. & For. 3 (1914) 162;
Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9 (1919) 128.
GUIGNARDIA MANIHOTI Sacc. var. DIMINUTA Sacc.
On Manihot utilissima'. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
18; Philip. Journ. Sci. 13 (1918) 168.
MASSALONGIELLA IMPERATAE Rehm.
On Imperata cylindrica. Rehm, Leafl. Philip. Bot. 8 (1916) 2956 —
Los Banos (Reyes, comm. Baker 3120).
MYCOSPHAERELLA ALOCASIAE Syd.
On Alocasia indica. Baker, Philip. Agr. & For. 3 (1914) 158; Ann.
Myc. 15 (1917) 205.
MYCOSPHAERELLA ARISTOLOCHIAE Syd.
Ann. Myc. 15 (1917) 205; Philip. Journ. Sci. 13 (1918) 377; Leafl.
Philip. Bot. 9 (1925) 3135.
MYCOSPHAERELLA BRIDELIAE Syd.
On Bridelia stipularis. Ann. Myc. 15 (1917) 206.
MYCOSPHAERELLA CARICAE Syd.
On Carica papaya. Baker, Philip. Agr. & For. 3 (1914) 159; Philip.
Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 118.
MYCOSPHAERELLA MUSAE Speg.
On Musa cavendishii. Baker, Philip. Agr. & For. 3 (1914) 162; Ann.
Myc. 15 (1917) 206.
On Musa sapientwnv. Philip. Journ. Sci. 13 (1918) 168; Ann. Myc.
21 (1923) 100; Philip. Agr. Rev. 18 (1925) 582.
On Musa paradisiaca sapientum. Phytopath. 9 (1919) 129.
On Musa textilis. Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9
(1919) 129.
MYCOSPHAERELLA OCULATA Syd.
On Premna sp. Ann. Myc. 15 (1917) 206.
On Premna odorata. Philip. Journ. Sci. 13 (1918) 377.
46,3 Baker: Lower Fungi 505
MYCOSPHAERELLA PERICAMPYLI Syd.
On Pericampylus ineanus. Ann. Myc. 15 (1917) 206; 21 (1923) 99;
Leafl. Philip. Bot. 9 (1925) 3135.
MYCOSPHAERELLA REYESII Syd.
On Sapindus saponaria. Baker, Philip. Agr. & For. 4 (1914) 164;
Ann. Myc. 15 (1917) 207.
SPHAERULINA SMILACINCOLA Rehm.
Ann. Myc. 20 (1922) 70.
CLYPEOSPHAERIACE^E
ANTHOSTOMELLA ARECAE Rehm.
On Areca catechu. Rehm, Leafl. Philip. Bot. 8 (1916) 2938 — Los Ba-
fios (Baker 3068); Baker, Philip. Agr. & For. 5 (1916) 74; Philip.
Journ. Sci. 13 (1918) 165.
ANTHOSTOMELLA ARENGAE (Rac.) Rehm.
Raciborski, Alg. und Pilze Javas 3 (1900) 27 (Awrswaldia) ; Rehm,
Philip. Journ. Sci. 8 (1900) 395 (Auerswaldia decipiens) ; 399 (An-
thostomella mindorensis) ; Sydow and Theissen, Ann. Myc. 13
(1900) 390; Rehm, Leafl. Philip. Bot. 8 (1916) 2940; Ann. Myc. 16
(1918) 223, 224.
ANTHOSTOMELLA ATRON1TENS Rehm.
On Donax cannaeformis. Ann. Myc. 15 (1917) 209.
ANTHOSTOMELLA CALAMI Rehm.
On Calamus. Rehm, Leafl. Philip. Bot. 8 (1916) 2939— -Mount Maqui-
ling (Baker 8186, 3345; Reyes, comm. Baker 83U5).
ANTHOSTOMELLA CALOCARPA Syd.
On Pandanws sabutan. Ann. Myc. 15 (1917) 209.
ANTHOSTOMELLA COCOINA Syd.
On Cocos nucifera. Philip. Agr. & For. 4 (1914) 160<; Philip. Journ.
Sci. 13 (1918) 166; Phytopath. 9 (1919) 122.
ANTHOSTOMELLA CORYPHAE Rehm.
On Corypha elata. Rehm, Leafl. Philip. Bot. 8 (1916) 2940— Los Ba-
fios (Baker 267 %) ; Ann. Myc. 15 (1917) 209.
ANTHOSTOMELLA CORYPHAE Rehm f. MINUTISSIMA Rehm.
On Corypha elata. Rehm, Leafl. Philip. Bot. 8 (1916) 2940— Los Ba-
fios (Evaristo, comm. Baker 2572).
ANTHOSTOMELLA DONACINA Rehm. f. ARENGAE Rehm.
On Arenga. Rehm, Leafl. Philip. Bot. 8 (1916) 2940 — Los Bafios (Ba*
ker 1797, 3064).
ANTHOSTOMELLA EUMORPHA (Sacc. and Paoli) Rehm.
Saccardo and Paoli, Myc. Malacc. No. 89 (Anthostoma eumorphum).
On Schizostachyum. Rehm, Leafl. Philip. Bot. 8 (1916) 2940— Los
Bafios (Baker 2021b).
506 The Philippine Journal of Science 1931
ANTHOSTOMELLA GRANDISPORA Penz. and Sacc.
On Bambusa and Schizostachyum. Penzig and Saccardo, Malpighia
11 (1897) 392; Rehm, Leafl. Philip. Bot. 8 (1916) 2939— Los Banos
(Reyes, comm. Baker 1U25); (Baker 1954a).
ANTHOSTOMELLA LUCENS Sacc.
On Pandanus. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 19 —
Mount Banahao (Baker 8860).
On Pandanus radicans. Leafl. Philip. Bot. 9 (1925) 3135.
ANTHOSTOMELLA MICRASPIS (Berk.) Sacc. and Trav.
Berkeley, Journ. Bot. (1842) 156 (Sphaeria) ; Currey, Trans. Linn.
Soc. Lond. Bot. 20 (1859) 321 (Sphaeria) ; Saccardo and Traverso,
Syll. Fung. 19 (1910) 77; 22 (1913) 101.
On fallen limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2938— Mount Ma-
quiling (Baker 2908; Reyes, comm. Baker 4025).
ANTHOSTOMELLA MIRABILIS (B. and Br.) v. Hoehn. (Astrocystis mirabilis B. and Br.)
On Bambusa, Sydow, Philip. Journ. Sci. § C 8 (1913) 485 (A. disco-
phora) ; Rehm, Leafl. Philip. Bot. 8 (1916) 2939— Los Banos (Reyes,
comm. Baker 3055, 84S3, 3404, 3652); Ann. Myc. 15 (1917) 209.
ANTHOSTOMELLA PANDANI (Rehm) Sydow.
On Pandanus. Baker, Leafl. Philip. Bot. 7 (1914) 2453 (Auerswal-
dia); Theissen and Sydow, Ann. Myc. 13 (1915) 300; Rehm, Leafl.
Philip. Bot. 8 (1916) 2*939— Mount Banahao (Baker 2236).
ANTHOSTOMELLA UBERIFORMIS Rehm.
On dead trunk. Rehm, Leafl. Philip. Bot. 8 (1916) 2937— Mount Ma-
quiling (Baker 8^11).
ASTROSPHARIELLA FUSISPORA Syd.
On Bambusa blumeana. Ann. Myc. 15 (1917) 209.
CEUTHOCARPON DEPOKENSE Penz. and Sacc.
On Dracontomelum cumingianum. Penzig and Saccardo, Malpighia
9 (1897) 405; Rehm, Leafl. Philip. Bot. 8 (1916) 2953— Los Banos
(Raimundo, comm. Baker 2191a).
CEUTHOCARPON PUNCTIFORME Sacc.
On Sterculia. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 21 — Los
Banos (Baker 3893).
CEUTHOCARPON TALAUMAE Rehm.
On Talauma villariana. Rehm, Leafl. Philip. Bot. 8 (1916) 2953 —
Los Banos (Raimundo, comm. Baker 28 43) .
CLYPEOSPHAERIA BAKERIANA Rehm.
On Eugenia bataanensis and Grewia stylocarpa. Rehm, Leafl. Philip.
Bot. 8 (1916) 2948 — Mount Maquiling (Baker 3481a) ; (Baker
3465); Ann. Myc. 15 (1917) 209.
46, 3 Baker: Lower Fungi 507
DIDYMOSPHAERIA ANISOMERA Sacc.
On Andropogon sorghum (Sorghum vulgare, Holcus sorghum). SAC-
CARDO, Nuovo Giorn. Bot. Ital. 23 (1916) 20 — Los Banos (Baker
3800); Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919) 138.
DIDYMOSPHAERIA CAESPITULOSA Sacc.
On Premna cumingiana. Ann. Myc. 13 (1915) 127.
DIDYMOSPHAERIA INCONSPICUA Rehm.
On Premna odorata. Rehm, Leafl. Philip. Bot. 8 (1916) 2948 — Los
Banos (Baker 2110b).
DIDYMOSPHAERIA STRITULA Penz. and Sacc.
On Bambusa vulgaris, Calamus, and Schizostachyum sp. Rehm, Leafl.
Philip. Bot. 8 (1916) 2948— Los Banos (Reyes, comm. Baker 190S) ;
Mount Maquiling (Reyes, comm. Baker 331+1+, 331+5)) Ann. Myc. 15
(1917) 208.
LINOSPORA ELASTICAE Koord.
Koorders, Bot. Untersuch (1917) 193.
On Ficus. Rehm, Leafl. Philip. Bot. 8 (1916) 2954— Mount Maqui-
ling (Copeland, comm. Baker 3179a).
LINOSPORA PANDANI Rehm.
On Pandanus sabotan and P. utilissima. Rehm, Leafl. Philip. Bot. 8
(1916) 2954 — Los Banos (Reyes, comm. Baker 3045) ; Mount Bana-
hao (Baker 221+8).
LINOSPORA SERIATA (Syd.) Rehm.
On Bambusa blumeana. Sydow, Philip. Journ. Sci. 8 (1916) 272
(Ophiobolus) ; Rehm, Leafl. Philip. Bot. 8 (1916) 2954— Mount
Maquiling (Baker 31+17).
PLEOSPORACEiE
DIDYMELLA CARICAE Tassi.
On Carica papaya. Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9
(1919) 118.
DIDYMELLA EUTYPOIDES Rehm.
On Bambusa. Rehm, Leafl. Philip. Bot. 8 (1916) 2943 — Los Banos
(Reyes, comm. Baker 1915c).
DIDYMELLA LUSSONIENSIS Sacc.
On Dolichos unifiorus. Baker, Philip. Agr. & For. 3 (1914) 161;
Phytopath. 9 (1919) 132.
On Dolichos lablab. Philip. Journ. Sci. 13 (1918) 167.
DIDYMELLA ORCHNODES Rehm.
On Goniothalamus. Rehm, Leafl. Philip. Bot. 8 (1916) 2*943— Mount
Maquiling (Baker 3085a).
DIDYMELLA SERIATA Rehm.
On Schizostachyum. Rehm, Leafl. Philip. Bot. 8 (1916) 2943 — Los
Banos (Baker 1951+b).
508 The Philippine Journal of Science m\i
DIDYMOSPHAERIA CAESPITULOSA Sacc.
On Premna cumingiana. Saccardo, Ann. Myc. 13 (1915) 127 — Los
Baiios (Baker 274.6).
DIDYMOSPHAERIA STRIATULA Penz. and Sacc.
Rehm, Leafl. Philip. Bot. 6 (1914) 2223— (Phaodothis gigantochloae) ;
Baker, Leafl. Philip. Bot. 8 (1914) 2455; Theissen and Sydow,
Ann. Myc. 13 (1915) 185.
LEPTOSPHAERIA ORTHOGRAMMA (B. and C.) Sacc.
Berkeley and Curtis, Cent. N. Am. Fung. (1853) No. 922 (Sphae-
ria); Saccardo, Syll. Fung. 2 (1883) 60.
On Zea mays. Rehm, Leafl. Philip. Bot. 8 (1916) 2951— Los Baiios
(Raimundo, comm. Baker 1996); Baker, Philip. Agr. & For. 5
(1916) 78; Philip. Journ. Sci. 13 (1918) 170.
METASPHAERIA CORRUSCANS Rehm.
On Capparis horrida. Rehm, Leafl. Philip. Bot. 8 (1916) 2950— Los
Baiios (Baker 14,29b).
METASPHAERIA INCOMPLETA Rehm.
On Eugenia. Rehm, Leafl. Philip. Bot. 8 (1916) 2949— Mount Maqui-
ling (Baker 2936b).
OPHIOCHAETE BAKERIANA Sacc.
On Calamus. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 21 — Mount
Maquiling (Baker 8775).
PHYSALOSPORA AFFINIS Sacc.
On Theobroma cacao. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
18 — Los Baiios (Baker 8779) ; Baker, Philip. Agr. & For. 5 (1916)
77; Philip. Journ. Sci. 13 (1918) 169; Phytopath. 9 (1919) 138.
PHYSALOSPORA BAMBUSAE (Rabh.) Sacc.
On Bambusa. Baker, Philip. Agr. & For. 3 (1914) 159.
PHYSALOSPORA BAMBUSICOLA Rehm.
On Bambusa. Baker, Philip. Agr. & For. 3 (1914) 159.
PHYSALOSPORA DINOCHLOAE Rehm.
On Dinochloa. Rehm, Leafl. Philip. Bot. 8 (1916) 2937 — Mount Ma-
quiling (Baker 2189a).
PHYSALOSPORA GUIGNARDIOIDES Sacc.
On Canavalia gladdata. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
19— Los Baiios (Baker 3809) ; Baker Philip. Agr. & For. 5 (1916)
74; Philip. Journ. Sci. 13 (1918) 166.
On Phaseolus spp. Phytopath. 9 (1919) 132.
PHYSALOSPORA HOYAE v. Hoehn.
V. Hoehnel, Kais. Ak. Wiess. Wien 114 (19Q7) 122.
On Hoya luzonica. Sydow, Leafl. Philip. Bot. 6 (1914) 2122 (P. ho-
yae) ; Rehm, Leafl. Philip. Bot. 8 (1916) 2937— Los Baiios (Baker
3093); Ann. Myc. 15 (1917) 207.
46, 3 Baker: Lower Fungi 509
PHYSALOSPORA PERIBAMBUSINA Rehm.
On Bambusa vulgaris. Rehm, Leafl. Philip. Bot. 8 (1916) 2937— Los
Bafios (Baker 6; Reyes, comm. Baker 1896, 1901).
TEPHROSTICTA FICINA Syd.
On Payena leeri. Ann. Myc. 15 (1917) 179.
On Coix lacryma-jobi. Ann. Myc. 15 (1917) 208.
OPHIOBOLUS HETEROSTROPHUS Drechsler.
Journ. Agr. Res. 31 (1925) 701-726; Philip. Agr. 19 (1931) 581-589.
OPHIOBOLUS NIPAE Henn.
On Nipa fructicans. Baker, Philip. Agr. & For. 3 (1914) 163.
OPHIOBOLUS ORYZAE I. Miyake.
Journ. Coll. Agr., Imp. Univ. Tokyo 2 (1910) 237-276.
OPHIOBOLUS ORYZINUS Sacc.
On Oryza sativa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 21 —
Los Bafios (Baker 877U, 3803 err. 3305) ; Baker, Philip. Agr. & Fbr.
5 (1916) 76; Reinking, Philip. Journ. Sci. 13 (1918) 168; Rein-
KING, Phytopath. 9 (1919) 131.
MASSARIACEJ3
MASSARIA BATAANENSIS Rehm.
On Eugenia bataanensis. Rehm, Leafl. Philip. Bot. 8 (1916) 2951 —
Mount Maquiling (Baker 3£81b).
MASSARINA RAIMUNDOI Rehm.
On Citrus nobilis. Baker, Philip. Agr. & For. 3 (1914) 160; Rein-
king, Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 119.
MASSARINULA BAMBUSICOLA Rehm.
On Bambusa vulgaris. Rehm, Leafl. Philip. Bot. 8 (1916) 2944 — Los
Bafios (Reyes, comm. Baker 1915b).
MASSARINULA DONACINA Rehm.
On Donax cannaeformis. Rehm, Leafl. Philip. Bot. 8 (1916) 2944 —
Los Bafios (Raimundo, comm. Baker 2013).
MASSARINULA OBLIQUA Sacc.
On Mischocarpus fuscescens. Saccardo, Ann. Myc. 13 (1915) 127 —
Los Bafios (Baker 2253).
MASSARINA RAIMUNDOI Rehm.
On Citrus nobilis. Reinking, Philip. Agr. 9 (1920-21) 133.
GNOMONIACE^E
GLOMERELLA CINGULATA (Stonem.) S. and r. S.
On Persea americana and Mangifera indica. Philip. Agr. 15 (1926)
128; Philip. Agr. Rev. 20 (1926) 271; 21 (1926) 81.
On Lagenaria leucantha. Philip. Agr. 14 (1926) 213.
510 The Philippine Journal of Science 1931
PHOMATOSPORA MIGRANS Rehm.
On Arenga saccharifera. Rehm, Leafl. Philip. Bot. 8 (1916) 2936
Los Banos {Reyes, comm. Baker 1455) ; Baker, Philip. tA.gr. & For.
5 (1916) 74; Ann. Myc. 16 (1918) 216.
VALSACEJE
DIAPORTHE CITRINCOLA Rehm.
On Citrus nobilis. Baker, Philip. Agr. & For. 3 (1914) 160; Philip.
Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 119; Reinking,
Philip. Agr. 9 (1920-21) 133.
DIAPORTHE RECONDITA Sacc.
On Gliricidia maculata. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
22— Los Banos (Baker 3793).
ENDOXYLA MANGIFERAE Henn.
On Mangifera indica. Baker, Philip. Agr. & For. 3 (1914) 162;
Philip. Journ. Sci. 13 (1918) 167; Phytopath. 9 (1919) 127.
EUTYPA BAMBUSINA Penz. and Sacc.
On Bambusa blumeana. Baker, Philip. A,gr. & For. 3 (1914) 159.
On dead culms of bamboo. Philip. Journ. Sci. 12 (1917) 377.
On Bambusa and Schizostachyum. Ann. Myc. 15 (1917) 213.
On Schizostachyum lumampao. Ann. Myc. 21 (1923) 101.
On Bambusa sp. Ann. Myc. 26 (1928) 431.
EUTYPA HETERACANTHA Sacc.
Syll. Fung. 1: 177; 9: 466.
On Citrus decumana. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
22— Los Banos (Baker 3897) ; Phytopath. 9 (1919) 119.
On Citrus maxima. Reinking, Philip. Agr. 9 (1920-21) 134.
EUTYPA LUDIBUNDA Sacc.
On branches. Ann. Myc. 15 (1917) 213.
PERONEUTYPELLA ARECAE Sydow.
On Areca catechu. Philip. Agr. & For. 4 (1914) 158; Reinking,
Philip. Journ. Sci. 13 (1918) 165.
On Cocos nucifera. Ann. Myc. 15 (1917) 213; Philip. Journ. Sci. 13
(1918) 166; Phytopath. 9 (1919) 122.
PERONEUTYPELLA GRAPHIDIOIDES Syd.
On Terminalia catappa. Phytopath. 9 (1919) 138.
EUTYPELLA CITRICOLA Speg.
On Citrus nobilis. Baker, Philip. Agr. & For. 3 (1914) 160; SACCAR-
DO, Nuovo Giorn. Bot. Ital. 23 (1916) 22 — Los Baiios (Baker 3898).
On Citrus maxima. Philip. Journ. Sci. 13 (1918) 165; 166; Phyto-
path. 9 (1919) 119; Reinking, Philip. Agr. 9 (1920-21) 133, 134.
On Citrus aurantifolia. Ann. Myc. 21 (1923) 101.
EUTYPELLA COCOS Ferd. and Winge.
On Cocos nucifera. Philip. Agr. & For. 4 (1914) 160; Philip. Journ.
Sci. 13 (1918) 166; Phytopath. 9 (1919) 122.
46,3 Baker: Lower Fungi 511
EUTYPELLA LEUCAENAE Rehro.
On Leucaena glauca. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
22 — Los Bafios (Baker 871*1).
EUTYPELLA LINEOLATA Rehm.
On Mallotus philippinensis. Rehm, Leafl. Philip. Bot. 8 (1916) 2955 —
Los Bafios (Baker 3060b).
EUTYPELLA MALLOTI Rehm.
On Mallotus philippinensis. Rehm, Leafl. Philip. Bot. 8 (1916) 2955 —
Los Bafios (Baker 3060a),
EUTYPELLA REHMIANA (Henn. and Nym.) v. Hohnel.
On Areca catechu. Philip. Journ. Sci. 13 (1918) 165.
THYRIDARIA CALAMINCOLA Rehm.
On Calamus. Rehm, Leafl. Philip. Bot. 8 (1916) 2957— Mount Ma-
quiling (Baker 3230b).
THYRIDARIA EMINENS Rehm.
On Streblus asper. Rehm, Leafl. Philip. Bot. 8 (1916) 2957 — Los Ba-
fios (Raimundo, comm. Baker 2977).
THYRIDARIA TARDA Bancroft.
On Theobroma cacao. Philip. Agr. & For. 4 (1915) 164; Phytopath.
9 (1919) 138.
MELANCONIDACE^E
VALSARIA CITRI Rehm.
On Citrus nobilis. Baker, Philip. Agr. & For. 3 (1914) 160; Rein-
king, Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 119;
Philip. Agr. 9 (1920-21) 133.
VALSARIA INSITIVA (de Not) Ces. and de Not.
On Morus alba. Baker, Philip. Agr. & For. 3 (1914) 162; Reinking,
Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9 (1919) 128.
DIATRYPACE^S
DIATRYPELLA BARLERIAE Syd.
On Barleria cristata. Baker, Philip. Agr. & For. 5 (1916) 74 — Los
Bafios.
DIATRYPELLA PSIDII Syd.
On Psidium guajava. Reinking, Phytopath. 9 (1919) 133.
MELOGRAMMATACEiE
BOTRYOSPHAERIA MINUSCULA Sacc.
On Thccbro-ina cacao. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
18— Los Banos (Baker 3777 y 3780) ; BAKER, Philip. Agr. & For. 5
(1916) 77; Reinking, Philip. Journ. Sci. 13 (1918) 169; Phytopath.
9 (1919) 139.
512 The Philippine Journal of Science i93i
XYLARIACEJE
HYPOXYLON ANNULATUM (Schw.) Mont.
On dead bark. Rehm, Leafl. Philip. Bot. 8 (1916) 2957 — Los Baiios
(Baker 2906) ; Ann. Myc. 15 (1917) 211.
HYPOXYLON ATROPURPUREUM Fr.
On Citrus nobilis. Philip. Journ. Sci. 13 (1918) 166.
On coccids. Reinking, Phytopath. 9 (1919) 119; Philip. Agr. 9
,(1920-21) 133.
HYPOXYLON CULMORUM Cke.
On Schizostachyum sp. Ann. Myc. 15 (1917) 212.
HYPOXYLON EFFUSUM Nitsch.
On bark of dead trees in the forest. Philip. Journ. Sci. 12 (1917)
378.
HYPOXYLON FREYCINETIAE Rehm.
On Freycinetia. Rehm, Leafl. Philip. Bot. 8 (1916) 2959— Mount Ma-
quiling (Baker 3J>16) ; Ann. Myc. 15 (1917) 211; 21 (1923) 101.
HYPOXYLON GRANULOSUM Bull.
Buluard, Champ. (1791) 176.
On dead branches. Rehm, Leafl. Philip. Bot. 8 (1916) 2958-— Los Ba-
iios (Reyes, comm. Baker 28S8).
HYPOXYLON HAEMATOSTROMA Mont.
On Schizostachyum and Bambusa. Rehm, Leafl. Philip. Bot. 8 (1916)
2958 — Mount Maquiling (Baker 390 U) ; (Reyes, comm. Baker 189 Ua).
HYPOXYLON MARGINATUM (Schw.) Berk.
On dead limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2958— Mount Ma-
quiling (Baker 34.83).
On wood. Ann. Myc. 15 (1917) 211.
On bark of dead trees in the forest. Philip. Journ. Sci. 12 (1917)
378.
HYPOXYLON MARGINATUM (Schw.) Berk. var. MAMMIFORME Rehm.
On fallen limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2958— Mount Ma-
quiling (Baker 3038).
HYPOXYLON RUBIGINEO-AREOLATUM Rehm var. MICROSPORUM Theiss.
Theissen, Ann. Myc. 6: 345.
On Polyscias nodosa. Rehm, Leafl. Philip. Bot. 8 (1916) 2958 — Mount
Maquiling (Baker 289k)*
On dead stems. Ann. Myc. 15 (1917) 212.
HYPOXYLON SUBEFFUSUM Spe«.
Spegazzini, Fung. Gnar. Pug. 1: No. 204; Sacoardo, Syll. Fung. 9
(1891) 556.
On rotten limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2*958— Los Bafios
(Reye,s, comm. Baker 2837); Ann. Myc. 15 (1917) 217.
46, 3 Baker: Lower Fungi 513
KRETZMARIA GHOMPHOIDEA Penz. and Sacc.
On rotten wood in forests. Philip. Journ. Sci. 12 (1917) 379.
NUMMULARIA CITRINCOLA Rehm.
On Citrus. Rehm, Leafl. Philip. Bot. 8 (1916) 2961— Los Banos (Ba-
ker 3062) ; Reinking, Philip. Agr. 9 (1920-21) 134.
NUMMULARIA FRAGILLIMA Rehm.
On Calamus. Rehm, Leafl. Philip. Bot. 8 (1916) 2959— Mount Ma-
quiling (Baker 8187).
NUMMULARIA GLYCYRRHIZA (B. and C.) Sacc.
On dead trunk. Ann. Myc. 15 (1917) 212.
NUMMULARIA LIANAE Rehm.
On a liana, perhaps Bauhinia. Rehm, Leafl. Philip. Bot. 8 (1916)
2959 — Mount Maquiling (Baker 2881 ) .
NUMMULARIA MEMORABILIS Rehm.
On dead wood. Rehm, Leafl. Philip. Bot. 8 (1916) 2960— Mount Ma-
quiling (Baker 81*32).
NUMMULARIA PAPYRACEA Rehm.
On dead trunk. Ann. Myc. 15 (1917) 212.
NUMMULARIA REYESIANA Rehm.
On Bambusa sp. and B. blumeana. Rehm, Leafl. Philip. Bot. 8 (1916)
2960 — Los Banos (Reyes, comm. Baker 1906) ; (Baker 1114., 162U,
257 U) ; Rehm, Leafl. Philip. Bot. 6: 2203 — Hypoxylon culmorum, non
Cke.).
On dead stems of bamboo. Philip. Journ Sci. 12 (1917) 378.
NUMMULARIA SCUTATA B. and C.
On fallen limbs and on Cyrilla. Rehm, Leafl. Philip. Bot. 8 (1916)
2961— Mount Maquiling (Baker 81*19, 31*31); (Baker 31*11*).
NUMMULARIA URCEOLATA Rehm.
On bark. Ann. Myc. 15 (1917) 212.
XYLARIACEiE
DALDINIA CONCENTRICA (Bolt.) Ces. and de Not.
On dead logs. Philip. Journ. Sci. 13 (1918) 378.
On Citrus maxima. Reinking, Philip. Agr. 9 (1920-21) 134.
On trunks of trees. Ann. Myc. 15 (1917) 212.
DALDINIA CONCENTRICA var. MICROSPORA (Starb.) Theiss.
On trunks of trees. Ann. Myc. 15 (1917) 212.
DALDINIA ESCHOLZII Ehr.
On trunks of trees. Ann. Myc. 15 (1917) 212.
XYLARIA ALLANTOIDEA Berk.
Ann. Myc. 15 (1917) 213.
263774 14
514 The Philippine Journal of Science 1931
XYLARIA CASTOREA Berk.
Rbinking, Philip. Agr. 9 (1920-21) 133.
XYLARIA CORNIFORMIS Fr.
On rotten logs. Philip. Journ. Sci. 12 (1917) 379.
XYLARIA EUGLOSSIA Fr.
On rotten logs. Ann. Myc. 15 (1917) 213.
XYLARIA GRAMMICA Mont.
On logs. Ann. Myc. 15 (1917) 213.
XYLARIA HYPOXYLON (L.) Grev. f. TROPICA Syd.
On rotting logs. Ann. Myc. 15 (1917) 212.
XYLARIA LUZONENSIS Henn.
On dead pods of Bauhinia lying on the ground in dense forests.
Philip. Journ. Sci. 12 (1917) 379.
XYLARIA NIGRIPES (Klot.) Sacc.
On deserted termite nests. Philip. Journ. Sci. 13 (1918) 227.
XYLARIA OBVATA Berk.
On logs. Ann. Myc. 15 (1917) 213.
XYLARIA PLEBEJA Ces.
On bark. Ann. Myc. 15 (1917) 213.
XYLARIA TABACINA (Kickx.) Berk.
On dead limbs. Rehm, Leafl. Philip. Bot. 8 (1916) 2961— Mount Ma-
quiling (Baker 3395).
XYLARIA TUBEROSA (Pers.) Cke.
On rotting wood and logs. Ann. Myc. 15 (1917) 213.
HYSTERIALES
HYPODERMATACEJE
LOPHODERMIUM ALEURITIS Rehm.
On dead leaves. Rehm, Leafl. Philip. Bot. 8 (1915) 2925 — Los Banos
(Baker SU4).
LOPHODERMIUM ARUNDINACEUM (Schrad.) Chev.
Schrader, Journ. f. d. Bot. 2 (1799) 62 (Hysterium) ; Fries, Syst.
Myc. 2 (1821) 590 (Hysterium); Chevalier, Flor. par. 1 (1826)
435; Saccardo, Syll. Fung. 2 (1883) 795.
On dead leaves of Livistona. Rehm, Leafl. Philip. Bot. 8 (1915) 2925 —
Mount Maquiling (Baker 34-22).
LOPHODERMIUM ARUNDINACEUM (Schrad.) Chev. f. VULGARE Fckl.
On dead Miscanthus japonicus. Rehm, Leafl. Philip. Bot. 8 (1915)
2926— Mount Maquiling (Baker 3527, 3540).
LOPHODERMIUM PASSIFLORAE Rehm.
Baker, Philip. Agr. & For. 3 (1914) 163.
46'3 Baker: Lower Fungi 515
LOPHODERMIUM PLANCHONIAE Rehm.
On dead leaves of Planchonia spectabilis. Rehm, Leafl. Philip. Bot. 8
(1915) 2925— Los Banos (Baker SO 80).
LOPHODERMIUM ROTUNDATUM Syd.
On Canarium sp. Ann. Myc. 15 (1917) 251.
HYSTERIACEJE
ALDONA STELLA NIGRA Rac.
On Pterocarpus sp. Leafl. Philip. Bot. 9 (1925) 3137.
HYSTERIUM ANCEPS Sacc.
On Streblus asper. Saccardo, Nuovo Giorn. Bot Ital. 23 (1916) 24 —
Los Banos (Baker 3831).
SCHIZOTHYRIUM ACERIS (P. Henn. and Lind.) Pat.
On Acer sp. Ann. Myc. 15 (1917) 251.
On Acer niveum. Ann. Myc. 21 (1923) 104.
PEZIZALES
CENANGIACE^E
CENANGIUM RLUMEANUM Rehm.
On dead Bambusa blumeana. Rehm, Leafl. Philip. Bot. 8 (1915)
2927 — Los Banos (Raimundo, comm. Baker 2927b).
PATELLARIACEJE
LAGERHEIMA DERMATOIDEA Rehm.
On dead Derris philippinensis. Rehm, Leafl. Philip. Bot. 8 (1915)
2928—Los Banos (Baker 2006a).
PACHYPATELLA ALSOPHILAE (Rac.) Theiss. and Syd.
On Alsophila. Raciborski, Paras. Alg. und Pilze Javas 2 (1900) 22
(Hysterostomella) — Java; Sydow, Philip. Journ. Sci. § C 8 (1913)
495 (Discodothis lobata Syd.) ; Baker, Leafl. Philip. Bot. 6 (1914)
2102 (Discodothis lobata).
On Cyathea caudata. Theissen and Sydow, Ann. Myc. 13 (1915)
228; 15 (1917) 252.
BULGARIACE.E
BULGARIASTRUM CAESPITOSUM Syd.
On Capparis sepiaria. Philip. Journ. Sci. 13 (1918) 361.
MOLLISIACE^
CALOPEZIZA MIRABILIS Syd.
On Premna odorata. Ann. Myc. 15 (1917) 218.
MOLLISIA RAVIDA Sydow.
On Lagerstroemia indica and L. speciosa. Philip. Agr. & For. 4 (1914)
161.
NIPTERA GREWIAE Rehm.
On leaves of Grewia. Rehm, Leafl. Philip. Bot. 8 (1915) 2928—Los
Banos (Baker 2885).
516 The Philippine Journal of Science i»si
TRICHOBELONIUM MELIOLOIDES Rehm.
On leaves of Gigantochloa scribneriana. Rehm, Leafl. Philip. Bot. 8
(1915) 2929— Hills back of Paete, Luzon (Baker 3115).
HELOTIACEJE
SCLEROTINIA NERVISEQUIA Schroet. v. BAMBUSACEA Rehm.
On dead Bambusa vulgaris and on dead leaves of Dimerocalyx longi-
pes. Rehm, Leafl. Philip. Bot. 8 (1915) 2930— Los Banos (Reyes,
comm. Baker 1911); Mount Maquiling (Reyes, comm. Baker 4119,
err. 8221).
PEZIZACE,E
HUMARIA CABALLINA Rehm.
On horse dung. Rehm, Leafl. Philip. Bot. 8 (1915) 2930— Mount Ma-
quiling (Copeland, comm. Baker 3637).
LACHNEA LIVID A (Schum.) Gill.
Saccardo, Syll. Fung. 8: 187.
On decaying plant remains on ground. Saccardo, Nuovo Giorn. Bot.
23 (1916) 24— Los Banos (Baker 2896, err. 3897).
LACHNEA LURIDA P. Henn. and E. Nym.
On Polyporus. Ann. Myc. 15 (1917) 252'.
PEZIZELLA OMBROPHILACEA Rehm.
On leaves of Psidium guajava. Rehm, Leafl. Philip. Bot. 8 (1915)
2929 — Los Banos (Raimundo, comm. Baker 198 4) ; Baker, Philip.
Agr. & For. 5 (1916) 76; Phytopath. 9 (1919) 133.
PILOCRATERA TRICHOLOMA (Mont.) P. Henn.
On logs. Ann. Myc. 15 (1917) 252.
PLICARIA BANANINCOLA Rehm.
On Musa sapientum. Philip. Journ. Sci. 13 (1918) 168.
On Musa paradisiaca sapientum. Phytopath. 9 (1919) 129.
PLICARIA TROPICA Rehm.
On burnt Bambusa. Rehm, Leafl. Philip. Bot. 8 (1915) 2931— Los
Banos (Raimundo, comm. Baker 1£45).
TRIBLIDIACEiE
TRYBLIDIELLA MINDANAENSIS P. Henn.
On branches. Ann. Myc. 15 (1917) 251.
On Premna. Philip. Journ. Sci. 12 (1917) 362.
On Hevea brasiliensis. Philip. Journ. Sci. 13 (1918) 167, 362.
On Citrus nobilis. Philip. Journ. Sci. 13 (1918) 166; Reinking, Philip.
Agr. 9 (1920-21) 133.
On Aberia gardneri. Ann. Myc. 21 (1923) 104.
TRYBLIDIELLA RUFULA (Spreng.) Saec.
On Citrus nobilis. Philip. Journ. Sci. 13 (1918) 166; Reinking, Philip.
Agr. 9 (1920-21) 133.
46, 3 Baker: Lower Fungi 517
PHACIDIALES
STICTIDACEJE
BRIARDIA MAQUILINGIANA Rehm.
On Tetrastigma. Rehm, Leafi. Philip. Bot. 8 (1915) 2927— Mount Ma-
quiling {Reyes, comm, Baker 38 20).
PROPOLIDIOPSIS ARENGA Rehm.
On Arenga. Rehm, Leafl. Philip. Bot. 8 (1915) 2927— Los Bafios
(Baker 2899).
PHACIDIACEJE
COCCOMYCES DUBIUS Rehm.
On Ficus minahassae. Rehm, Leafl. Philip. Bot. 8 (1915) 2926 — Los
Bafios (Reyes, comm. Baker 3480).
COCCOMYCES QUADRATUS (Schw. and Kze.) Karst. var. PHILIPPINUS Rehm.
On dead leaves of Neolitsea. Rehm, Leafl. Philip. Bot. 8 (1915) 2926 —
Mount Maquiling (Baker 8W).
RHAGADOLOBIUM BAKERIANUM Sacc.
On Cyathus. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 24 — Mount
Maquiling (Baker 38U) ; Ann. Myc. 20 (1922) 73.
RHYTISMA LAGERSTROEMIA Rabh.
Rabenhorst, Hedw. 31 (1878) ; Berkeley and Broome, Grev. 6 (1878)
110 (R. pongamiae).
On Lagerstroemia indica and L. speciosa. Philip. Agr. & For. 4
(1914) 161; Rehm, Leafl. Philip. Bot. 8 (1915) 2926— Morong Valley,
Rizal Province (Raimundo, comm. Baker 2580) ; Philip. Journ. Sci.
12 (1917) 362; Ann. Myc. 15 (1917) 251.
MYRIANGIALES
ELSINOEJ3
ELSINOE CANAVALIAE Rac.
On Canavalia ensiformis. Baker, Philip. Agr. & For. 3 (1914) 159;
Ann. Myc. 15 (1917) 255.
On Canavalia gladiata. Philip. Journ. Sci. 13 (1918) 165.
On Phaseolus spp. Phytopath. 9 (1919) 132.
MYRIANGIUM DURIAEI Mont.
On coccids. Reinking, Philip. Agr. 9 (1920-21) 133, 146.
PHYCOMYCETES
OOMYCETES
CHYTRIDIALES
SYNCHYTRIACE^D
WORONINELLA AECIDIOIDES (Peck.) Syd.
Peck, 24th Rep. N. Y. State Mus. 88 (1872) (Uredo) ; Thuembn, Myc.
Univ. No. 538 (1876) (Uredo peckii) ; Farlow, Bull. Bussey Inst. 2
518 The Philippine Journal of Science 1931
(1878) 229 (Synchytrium fulgens v. decipiens) ; Farlow, Bot. Gaz.
10 (1885) 240 (Synchytrium decipiens); Peck, in C. L. Shear, N.
Y. Fungi. Exsicc. No. 126 (1895) (Synchytrium aecidioides) ; Wil-
son and Seaver, Ascom. & Lower Fungi. Exsicc. No. 72 (Synchy-
trium aecidioides) (1909) ; Wilson and Seaver, Mycologia 1 (1909)
272 (Synchytrium aecidioides) ; Baker, Leafl. Philip. Bot. 6 (1914)
2149 (Synchytrium aecidioides); Sydow, Ann. Myc. 12 (1914) 485.
WORONINELLA DOLICHI (Cke.) Syd.
Cooke, Grevilea 10 (1882) 127 (Aecidium) ; Hennings, Engl. Bot.
Jahrb, 38 (1905) 103 (Uromyces vignicola) ; Sydow, Ann. Myc. 12
(1914) 486 — On Dolichos gibbosus, Glycine javanica, Dunbaria fer-
rignes, and Vigna sinensis in Ant. Africa, South Africa, India,
and Philippines.
On Dolichos lablab. Philip. Journ. Sci. 13 (1918) 167.
WORONINELLA PSOPHOCARPI Rac.
Raciborski, Zeitschr. f. Pflanzenk 195 (1898) ; Sydow, Ann. Myc. 1
(1903) 15 (Uromyces); 13 (1914) 486 — On Psophocarpus in Java,
Philippines, and West Africa.
On Psophocarpus tetragonolobus. Baker, Philip. Agr. & For. 5 (1916)
76; Philip. Journ. Sci. 13 (1918) 169; Phytopath. 9 (1919) 133;
Ann. Myc. 26 (1928) 414.
WORONINELLA PUERARIAE (Henn.) Syd.
Hennings, Engl. Bot. Jahrb. 15 (1892) 6 (Aecidium); Dietel, Engl.
Bot. Jahrb. 28 (1900) 282 (Uromyces); Miyabe, Bot. Mag. Tokyo
19 (1905) 199 (Synchytrium); Sydow, Ann. Myc. 12 (1914) 486—
On Pueraria in Java, New Guinea, Philippines, and Japan.
MYCOCHYTRIDIACEiE
AMPHOROMORPHA ENTOMOPHILA Thaxter.
Thaxter, Bot. Gaz. 58 (1914) 251 — Manila, on Diochus conicicallis
Mots, and on Labia sp. (Banks).
PYTHIACE.E
PYTHIUM DEBARYANUM Hesse.
Hesse, Phytium de Baryanum (1874) 34; Sadebeck, Stiz. Bot. ver.
Brandeb. (1874) 116 (P. esquiseti) ; Lohde, Uebe in paras Pilze
(1874) 203 (Lucidium pythiodes) ; Smith, Gard. Chron. 5 (1876)
656; Sadebeck, Tagebl. 49 Vers, deutsch. Naturf. u. Aerzte
(1876) (P. autumnale) ; Berlese and de Toni, Syll. Fung. 7 (1888)
271 (excl. syn. P. vexans) ; Atkinson, Bull. Cornell Exp. Sta. 94
(1895) (Artotrogus) ; Butler, Mem. Dept. Agr. India 1 (1907) No.
5, 86.
On Camelia sativa, Lepidium sativum, and Ricinus communis. Philip.
Agr. & For. 5 (1916) 70.
On Carica papaya, Lycopersicum e?culentum, and Nicotiana tabacum.
Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 30.
On Oryza sativa. Philip. Agr. 15 (1926) 290, 362.
46' 3 Baker: Lower Fungi 519
PERONOSPORALES
PERONOSPORACEiE
PHYTOPHTHORA COLOCASIAE Rac.
On Colocasia esculentum (Colocasia antiquorum). Raciborski, Paras
Alg. Pilze Javas 1 (1900) 9— Java; Sydow and Butler, Ann. Myc.
5 (1907) 512; Butler and Kulkarni, Mem. Dept. Agr. India 5
(1913) No. 5, 233-259; Mendiola and Esfino, Philip. Agr. & For.
5 (1916) 68— Los Baiios; Baker, Philip. Agr. & For. 5 (1917) 74;
Reinking, Philip. Journ. Sci. 13 (1918) 167; Reinking, Phytopath.
9 (1919) 123; Philip. Agr. Rev. 18 (1925) 560; Philip. Agr. 14
(1925-26) 439.
PHYTOPHTHORA FABERI Maubl.
Maublanc, L'Agr. Prat. Pays Chauds No. 79 (1909) 315; Coleman,
Ann. Myc. 8 (1910) 621 (P. theobromae) .
On Theobroma, Hevea, and Artocarpus. Saccardo and Trotter, Syll.
Fung. 21 (1912) 86.
On Theobroma cacao and Carica papaya. Mendiola and Espino,
Philip. Agr. & For. 5 (1916) 66— Los Bafios; Baker, Philip. Agr. &
For. 5 (1916) 77; Reinking, Philip. Journ. Sci. 13 (1918) 166.
On Cocos nucifera. Reinking, Philip. Journ. Sci. 14 (1919) 131;
Journ. Agr. Res. 25 (1923) 267.
On Citrus spp. Ocfemia and Roldan, Am. Journ. Bot. 14 (1927) 1.
PHYTOPHTHORA INFESTANS (Mont.) de Bary.
On Ly coper sicum esculentum. Phytopath. 9 (1919) 127.
On Solarium tuberosum. Philip. Agr. & For. 5 (1916) 65; Philip.
Journ. Sci. 13 (1918) 169, 361; Philip. Agr. 10 (1922) 348.
PHYTOPHTHORA MELONGENAE K. Sawada.
On Solanum melingena. Noji Shikenjo Tokubetsu Hokoku 2 (1915)
77-79; Mycologia 9 (1917) 249-253; Philip. Agr. 14 (1925) 317-328.
PHYTOPHTHORA PHASEOLI Thaxter.
On Sandoricum koetjape (S. indicum). Clara, Philip. Journ. Sci. 35
(1928) 423.
SCLEROSPORA PHILIPPINENSIS Weston and SCHLEROSPORA SPONTANEA Weston.
(Sclerospora maydis (Rac.) Butler.)
On Zea mays. Raciborski, Ber. de Deutsch. Bot. Gessellsch. 15 (1897)
475 (Peronospora); Saccardo and Sydow, Syll. Fung. 14 (1899)
460 (Peronospora) ; Berlese, Riv. Pat. Veg. 10 (1904) 219 (Peronos-
pora) ; Butler, Mem. Dept. Agr. Ind. Bot. 5 (1913) No. 5, 275; Bak-
er, Philip. Agr. & For. 5 (1916) 78— Los Banos; Philip. Journ. Sci.
13 (1918) 131; Phytopath. 9 (1919) 139; Journ. Agr. Res. 19
(1920) 97; Philip. Agr. 8 (1920) 333; Journ. Agr. Res. 20 (1921)
678; Phytopath. 11 (1921) 372; Journ. Agr. Res. 20 (1921) 559;
23: 276, 726; Philip. Agr. 15 (1926) 127.
SCLEROSPORA SACCHARI Miyake.
On Saccharum officinarum. Phytopath. 11 (1921) 371.
520 The Philippine Journal of Science mi
ZYGOMYCETES
MYCORALES
MUCORACEJE
RHIZOPUS ARTOCARPI Rac.
On Artocarpus Integra (Artocarpus integrifolia). Baker, Philip. Agr.
& For. 3 (1914) 158; Philip. Journ. Sci. 13 (1918) 361; Rbinking,
Philip. Journ. Sci. 13 (1918) 131; Philip. Agr. 12 (1923-24) 465.
On Artocarpus communis. Reinking, Philip. Journ. Sci. 13 (1918)
131; Phytopath. 9 (1919) 116.
On Artocarpus incisa. Reinking, Phytopath. 9 (1919) 116.
RHIZOPUS NIGRICANS Ehrenberg.
On fiber of Musa textilis. Philip. Journ. Sci. 32 (1927) 79.
PILOBOLACE^E
PILOBOLUS LENTIGER Cda.
Corda, Icon. Fung. 1 (1837) 22; Saccardo, Syll. Fung. 7 (1837) 188,
Grove, Journ. Bot. (1884) 132 (P. kleinii var. sphaerospora) .
On horse dung. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 25— -Los
Banos (Baker 8892).
FUNGI IMPERFECT!
SPHAERIOPSIDALES
SPHAERIOIDACEJE
ASTEROMA PHASEOLI Brun.
Saccardo, Syll. Fung. 10 (1916) 219.
On pods of Phaseolus vulgaris. Saccardo, Nuovo Giorn. Bot. Ital.
23 (1916) 25— Los Banos (Baker 3728) ; Baker, Philip. Agr. & For.
5 (1916) 76; Reinking, Philip. Journ. Sci. 13 (1916) 166; Phyto-
path. 9 (1919) 132.
BAKEROPHOMA SACCHARI Diedicke.
On Saccharum officinarum. Baker, Philip. Agr. & For. 5 (1916) 76
Los Banos; Diedicke, Ann. Myc. 14 (1916) 62; Reinking, Philip.
Journ. Sci. 13 (1918) 166; Philip. Agr. Rev. 11 (1918) 275; Phyto-
path. 9 (1919) 134; Philip. Agr. Rev. 14 (1921) 430.
BOTRYODIPLODIA ANCEPS Sacc. and Syd.
On Morus alba. Baker, Philip. Agr. & For. 3 (1914) 162; Ann. Myc.
15 (1917) 28; Reinking, Philip. Journ. Sci. 13 (1918) 166; Phyto-
path. 9 (1919) 128.
BOTRYODIPLODIA CURTA Sacc.
On Ricinus communis. Ann. Myc. 15 (1917) 258.
CONIOTHYRIUM COFFEAE Heim.
On Coffea arabica. Baker, Philip. Agr. & For. 3 (1914) 160; Rein-
king, Phytopath. 9 (1919) 122.
On Coffea spp. Reinking, Philip. Journ. Sci. 13 (1918) 166.
46,3 Baker: Lower Fungi 521
CYTOSPORA ABERRANS Sacc.
On Citrus nobilis. Baker, Philip. Agr. & For. 3 (1914) 160; Rein-
king, Philip. Journ. Sci. 13 (1918) 166.
On Citrus sp. Ann. Myc. 15 (1917) 256.
On coccids. Reinking, Phytopath. 9 (1919) 119.
CYTOSPORA PALMICOLA B. and Cke.
On Cocos nucifera. Ann. Myc. 15 (1917) 256; Reinxing, Philip.
Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 122.
DIPLODIA ARTOCARPI Sacc.
Baker, Philip. Agr. & For. 3 (1914) 158.
On Artocarpus communis. Reinking, Philip. Journ. Sci. 13 (1918)
166.
On Artocarpus incisa. Reinking, Phytopath. 9 (1919) 116.
DIPLODIA ARTOCARPINA Sacc.
On Artocarpus integra (A. integrifolia). Baker, Philip. Agr. & For.
3 (1914) 158; Reinking, Philip. Journ. Sci. 13 (1918) 166; Phyto-
path. 9 (1919) 116.
DIPLODIA CARICAE Sacc.
On Carica papaya. Baker, Philip. Agr. & For. 3 (1914) 159; Ann.
Myc. 15 (1917) 257; Reinking, Philip. Journ. Sci. 13 (1918) 166;
Phytopath. 9 (1919) 118.
DIPLODIA CIRCINANS B. and Br.
On Yucca aloifolia. Ann. Myc. 15 (1917) 257.
DIPLODIA COCOCARPA Sacc.
On Cocos nucifera. Baker, Philip. Agr. & For. 3 (1914) 160; Rein-
king, Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 122.
DIPLODIA COCOCARPA var. MALACCENSIS Tassi.
On Cocos nucifera. Reinking, Philip. Journ. Sci. 13 (1918) 166.
DIPLODIA CREBRA Sacc.
On fruits of Musa sapientum. Saccardo, Nuovo Giorn. Bot. Ital. 23
(1916) 28— Los Banos (Baker 3743 f err. 3745) ; Baker, Philip. Agr.
& For. 5 (1916) 75; Reinking, Philip. Journ. Sci. 13 (1918) 168.
On Musa sp. Reinking, Phytopath. 9 (1919) 127.
DIPLODIA DATURAE Sacc.
On Datura alba. Ann. Myc. 15 (1917) 257.
DIPLODIA DURIONIS Sacc. and Syd.
On Durio zibethinus. Baker, Philip. Agr. & For. 3 (1914) 161.
DIPLODIA MANIHOTI Sacc.
On Manihot utilissima. Baker, Philip. Agr. & For. 3 (1914) 162;
Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 28 — Los Banos (Baker
3888); Reinking, Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9
(1919) 128.
522 The Philippine Journal of Science mi
DIPLODIA MORI West.
Saccardo, Syll. Fung. 3: 351.
On Mortis alba. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 28—
Los Banos (Baker 3818); Baker, Philip. Agr. & For. 5 (1916) 75-
Ann. Myc. 15 (1917) 257; Reinking, Philip. Journ. Sci. 13 (1918)
168; Phytopath. 9 (1919) 128.
DIPLODIA PHASEOLINA Sacc.
On Phaseolus lunatas. Baker, Philip. Agr. & For. 3 (1914) 163-
Ann. Myc. 15 (1917) 257.
On Phaseolus vulgaris. Baker, Philip. Agr. & For. 5 (1916) 76— Los
Banos; Reinking, Philip. Journ. Sci. 13 (1918) 169; Phytopath. 9
(1919) 132.
DIPLODIA RICINICOLA Sacc.
On Ricinus communis. Ann. Myc. 15 (1917) 257.
DIPLODIA SYNEDRELLAE Sacc.
On Synedrella nodiflora.
DIPLODINA DEGENERANS Diedicke.
On Solarium melongena. Baker, Philip. Agr. & For. 5 (1916) 77
Los Banos; Ann. Myc. 14 (1916) 64.
HAPLOSPORA MANILENSIS Sacc.
On Ricinus communis. Ann. Myc. 15 (1917) 2*57.
DOTHIORELLA CRASTOPHILA Sacc.
On Bambusa. Ann. Myc. 15 (1917) 257.
LASIODIPLODIA THEOBROMAE (Pat.) Griff, and Maubl.
On Theobroma cacao. Baker, Philip. Agr. & For. 3 (1914) 164;
4 (1915) 164; Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 28—
Los Banos (Baker 2729a, 2778) ; Philip. Agr. & For. 5 (1916) 77;
Ann. Myc. 15 (1917) 258; Reinking, Philip. Journ. Sci. 13 (1918)'
169; Phytopath. 9 (1919) 138; Philip. Agr. 8 (1920) 237.
On Ipomoea batatas. Baker, Philip. Agr. & For. 5 (1916) 77 Los
Banos.
On Carica papaya, Citrus maxima, and Dioscorea esculenta. Rein-
king, Philip. Journ. Sci. 13 (1918) 166.
On Hevea brasiliensis. Ann. Myc. 21 (1923) 105.
MACROPHOMA ARENGAE Sacc.
On Arenga saccharifera. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
27 — Los Banos (Baker 2827) .
MACROPHOMA CYANOPSIDIS Syd.
On Cyanopsis psoraleoides. Baker, Philip. Agr. & For. 3 (1914) 161.
MACROPHOMA MUSAE (Cke.) Berl. and Vogl. (Phoma musae Carpenter.)
On Musa sapientum. Baker, Philip. Agr. & For. 3 (1914) 162; Ann.
Myc. 15 (1917) 256; Philip. Agr. Rev. 14 (1921) 425; Phytopath."
46> 3 Baker: Lower Fungi 523
12 (1922) 101; Ann. Myc. 21 (1923) 105; Philip. Agr. Rev. 18
(1925) 582; Philip. Agr. 15 (1926) 469.
On Musa paradisiaca sapieyitum. Reinking, Phytopath. 9 (1919)
128.
On Musa textilis. Reinking, Philip. Journ. Sci. 13 (1918) 168.
MACROPHOMA OBSOLETA Sacc.
On Capparis horrida. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
26.
MACROPHOMA TRICHOSANTHIS Syd.
On Trichosanthes anguina. Baker, Philip. Agr. & For. 5 (1916)
77 — Los B alios.
On Cucumis sativum. Phytopath. 9 (1919) 124.
MICRODIPLODIA PASSERINIANA (Thiim.) Allesch.
Saccardo, Syll. Fung. 3: 371.
On Arenga saccharifera. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
28 — Los Banos (Baker 3866) ; Baker, Philip. Agr. & For. 5 (1916)
74.
APHYSA DESMODII Syd. (= Pazschkiella philippinensis Yates.)
On Desmodium sinuoswin. Ann. Myc. 15 (1917) 205; 20 (1922) 73;
21 (1923) 99; 26 (1928) 435.
On Dunbaria sp. Philip. Journ. Sci. 13 (1918) 380.
PHOMA BAKERIANA Sacc.
On Vigrm spp. Philip. Agr. & For. 4 (1914) 164; Reinking, Philip.
Journ. Sci. 13 (1918) 170; Phytopath. 9 (1919) 139.
PHOMA CITRICARPA McAlpine.
On Citrus spp. Philip. Journ. Sci. 17 (1920) 640.
PHOMOPSIS CALANTHES Sacc.
On Calanthes. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 27 —
Mount Maquiling (Baker 3821*).
PHOMOPSIS CAPSICI (Magnaghi) Sacc.
Saccardo, Syll. Fung. 18: 256.
On Capsicum annuum. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
27— Los Banos (Baker 3749) ; Baker, Philip. Agr. & For. 5 (1916)
74; Reinking, Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9
(1919) 117.
PHOMOPSIS CINERESCENS (Sacc.) Bubak.
On Ficus sp. Ann. Myc. 15 (1917) 256.
PHOMOPSIS DIOSCOREAE Sacc.
On Dioseorea esculenta. Reinking, Philip. Journ. Sci. 13 (1918)
167.
PHOMOPSIS GLIRICIDIAE Syd.
On Gliricidia maculata. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
27 — Los Banos (Baker 3820).
524 The Philippine Journal of Science 1931
PHOMOPSIS PALMICOLA (Wint.) Sacc. f. ARECAE Sacc.
On Areca catechu. Saccardo, Ann. Myc. 13 (1915) 128 — Los Baiios
(Raimwido, comm. Baker 2953) ; Baker, Philip. Agr. & For. 5
(1916) 73 — Los Bafios; Reinking, Philip. Journ. Sci. 13 (1918)
165.
PHOMA HERBARUM Westd.
On Manihot utilissima. Reinking, Philip. Journ. Sci. 13 (1918) 168.
PHOMA OLERACEA Sacc.
On Dioscorea spp. Baker, Philip. Agr. & For. 3 (1914) 161; Rein-
king, Phytopath. 9 (1919) 124.
On Dioscorea esculenta. Reinking, Philip. Journ. Sci. 13 (1918) 167.
PHOMA SABDARIFFAE Sacc.
On Hibiscus sabdariffa. Baker, Philip. Agr. & For. 4 (1914) 161;
Ann. Myc. 15 (1917) 256; Reinking, Philip. Journ. Sci. 13 (1918)
167; Phytopath. 9 (1919) 126.
PHOMA SESAMINA Sacc.
On Sesamum orientate (S. indicum). Baker, Philip. Agr. & For. 3
(1914) 164; Reinking, Philip. Journ. Sci. 13 (1918) 169; Phyto-
path. 9 (1919) 136.
PHOMA SOLANOPHILA Oud.
Saccardo, Syll. Fung. 16: 870.
On Solatium melongena. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
27 — Los Baiios (Baker 8825); Reinking, Philip. Journ. Sci. 13
(1918) 169.
PHELLOSTROMA HYPOXYLOIDES Syd.
On Areca catechu. Philip. Agr. & For. 4 (1914) 158; Philip. Journ.
Sci. 13 (1918) 165.
PHOMOPSIS ARECAE Syd.
On Areca catechu. Baker, Philip. Journ. Agr. & For. 4 (1914) 158;
Reinking, Philip. Journ. Sci. 13 (1918) 165.
PHYLLOSTICTA CIRCUMSEPTA Sacc.
On Citrus nobilis. Baker, Philip. Agr. & For. 3 (1914) 160.
On Citru^s maxima. Reinking, Philip. Journ. Sci. 13 (1918) 166.
On Citrus spp. Reinking, Phytopath. 9 (1919) 120; Philip. Agr. 9
(1920-21) 135.
PHYLLOSTICTA COCOPHYLLA Pass.
On Cocos nucifera. Reinking, Philip. Journ. Sci. 13 (1918) 167;
Phytopath. 9 (1919) 122.
PHYLLOSTICTA DENSISSIMA Sacc.
On Capparis horrida. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
26— Los Baiios (Baker 3787a).
PHYLLOSTICTA DYSOXYLI Sacc.
On Dysoxylum. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 26 —
Mount Maquiling (Baker 3795).
46> 3 Baker: Lower Fungi 525
PHYLLOSTICTA EUCHLAENAE Sacc.
On Euchlaena luxurians. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
25— Los Baiios (Baker 373b) ; Baker, Philip. Agr. & For. 5 (1916)
75.
PHYLLOSTICTA GLUMARUM Sacc.
On Oryza sativa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 25
Los Baiios (Baker 3871, err. 3371); Baker, Philip. Agr. & For. 5
(1916) 75; Ann. Myc. 15 (1917) 256; Reinking, Philip. Journ. Sci.
13 (1918) 168; Phytopath. 9 (1919) 131.
PHYLLOSTICTA GRAFFIANA Sacc.
On Dwscorea aculeata. Ann. Myc. 15 (1917) 255.
On Dioscorea esculenta. Reinking, Philip. Journ. Sci. 13 (1918)
167, 381.
PHYLLOSTICTA INSULARUM Sacc.
On Anona muricata. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
26 — Los Banos (Baker 3795) ; Baker, Philip. Agr. & For. 5 (1916)
73; Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9
(1919) 115.
PHYLLOSTICTA MANHOTICOLA Syd.
On Manihot dichotoma. Baker, Philip. Agr. & For. 3 (1914) 162;
Reinking, Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9 (1919)
127.
PHYLLOSTICTA MIURAI I. Miyake.
Saccardo, Syll. Fung. 22: 864.
On Oryza sativa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 26 —
Los Banos (Baker 3811); Baker, Philip. Agr. & For. 5 (1916) 75;
Philip. Journ. Sci. 13 (1918) 381; Reinking, Philip. Journ. Sci. 13
(1918) 168; Phytopath. 9 (1919) 131.
PLACOSPHAERIA DURIONIS Syd.
On Durio zibethinu-s. Baker, Philip. Agr. & For. 3 (1914) 161.
PLACOSPHAERIA TIGLII Henn.
On Croton tigliwm. Baker, Philip. Agr. & For. 4 (1914) 161; Ann.
Myc. 15 (1917) 256; Philip. Journ. Sci. 13 (1918) 381.
RHABDOSPORA SYNEDRELLAE Sacc.
On dead stems of Synedrella nodiflora. Saccardo, Ann. Myc. 13
(1915) 128— Los Banos (Baker 3228).
SEPTORIA PALMARUM Sacc.
On Corypha elata. Baker, Philip. Agr. & For. 3 (1914) 160.
SEPTOSPORIELLA PHILIPPINENSIS Sacc.
On Saccharum spontaneum. Saccardo, Syll. Fung. 3 (1916) 29—
Los Baiios (Baker 3742).
STAGONOSPORA VARIANS Sacc.
On Symploeum whitfordii. Ann. Myc. 15 (1917) 259.
526 The Philippine Journal of Science 1931
TRAVERSOA DOTHIORELLOIDES Sacc. and Syd.
On Morus alba. Baker, Philip. Agr. & For. 3 (1914) 162; Reinking,
Philip. Journ. Sci. 13 (1918) 166; Phytopath. 9 (1919) 128.
On Citrus nobilis. Ann. Myc. 15 (1917) 257.
TRAVERSOA EXCIPULOIDES Sacc.
Ann. Myc. 15 (1917) 257.
TRAVERSOA EXCIPULOIDES Sacc. and Syd. var. DISTANS Sacc. and Syd.
On Gliricidia septum. Ann. Myc. 15 (1917) 257.
VERMICULARIA BREVISETA Sacc.
On Synedrella nodiflora. Ann. Myc. 15 (1917) 267.
VERMICULARIA CAPSICI Syd.
On Capsicum annuum. Reinking, Philip. Journ. Sci. 13 (1918) 165;
Phytopath. 9 (1919) 117.
VERMICULARIA FALLAX Sacc.
On Passiflora quadrangular is. Baker, Philip. Agr. & For. 3 (1914)
163.
VERMICULARIA HORRIDULA Sacc.
On Dolichos uniflorus. Baker, Philip. Agr. & For. 3 (1914) 161;
Reinking, Phytopath. 9 (1919) 132.
On Dolichos lablab. Reinking, Philip. Journ. Sci. 13 (1918) 167.
VERMICULARIA MERRILLIANA Sacc.
On Datura alba. Ann. Myc. 15 (1917) 267.
VERMICULARIA SESAMINA Sacc.
On Sesamum orientale (S. indicum). Reinking, Philip. Journ. Sci.
13 (1918) 169; Phytopath. 9 (1919) 136.
VERMICULARIA XANTHOSOMATIS Sacc.
On Xanthosoma sagitti folium. Saccardo, Nuovo Giorn. Bot. Ital. 23
(1916) 28— Los Bafios (Baker 3750) ; Baker, Philip. Agr. & For.
5 (1916) 78; Reinking, Phytopath. 9 (1919) 139.
YPSILONIA CUSPIDATA Leveille.
On leaves on one of the Anonacese. Leveille, Ann. Sci. Nat. (1846)
284— Manila (Cuming); Saccardo, Syll. Fung. 3 (1884) 216.
On Cyclostemon sp. Ann. Myc. 15 (1917) 261.
NECTRIOIDACEJE
ASCHERSONIA CINNABARINA P. Henn.
On Astronia. Ann. Myc. 15 (1917) 261.
ASCHERSONIA CONFLUENS Henn.
Hennings, Monsunia 1 (1899) 37; Hedwigia 145 (1902) (A. phthurioi-
des); Baker, Leafl. Philip. Bot. 6 (1914) 2155; Petch, Ann. Roy.
Bot. Gard. Peradeniya 5 (1914) 526 (stage of Hypocrella mollii
Koord.).
46, 3 Baker: Lower Fungi 527
ASCHERSONIA LECANIOIDES P. Henn.
On Melastoma. Ann. Myc. 15 (1917) 261.
ASCHERSONIA PARAENSIS Henn.
Saccardo, Syll. Fung. 18: 413.
On coccids on Psidium guajava. Saccardo, Nuovo Giorn. Bot. Ital.
23 (1916) 29— Hills back of Paete, Laguna Province (Baker 3790) ;
Baker, Philip. Agr. & For. 5 (1916) 76; Reinking, Phytopath. 9
(1919) 133.
ASCHERSONIA PLACENTA B. and Br.
Berkeley and Broome, Journ. Linn. Soc. Bot. 14 (1873) 89; Hen-
NINGS, Engl. Bot. Jahrb. 25 (1898) 509 (A. novo-guineensis) ;
Penzig and Saccardo, Malpighia (1901) 236 (A. javanica) ; Hen-
nings, Hedwigia (1902) 145 (A. lecanioides) ; Baker, Leafl. Philip.
Bot. 6 (1914) 2155 (A. lecanioides and A. novoguineensis) ; Petch,
Ann. Roy. Bot. Gard. Peradeniya 5 (1914) 527.
ASCHERSONIA SAMOENSIS Henn.
Hennings, Engler's Bot. Jahrb. 23 (1896) 289; Monsunia 1 (1899)
37 (A. cinnabarina) ; Patouillard and Hariot, Bull. Soc. Myc. Fr.
20 (1904) 65 (A. napoleonae) ; Baker, Leafl. Philip. Bot. 6 (1914)
2154 (A. cinnabarina) ; Petch, Ann. Roy. Bot. Gard. Peradeniya 5
(1914) 526 [stage of Hypocrella discoidea (B. and Br.) Sacc.].
ASCHERSONIA SCLEROTOIDES Henn.
Hennings, Hedwigia (1902) 146; Patouillard, Bull. Soc. Myc. Fr.
22 (1906) 59 (A. pisiformis) ; Baker, Leafl. Philip. Bot. 7 (1914)
2514; Petch, Ann. Roy. Bot. Gard. Peradeniya 5 (1914) 525 (stage
of Hypocrella reineckiana Henn.).
On Citrus sp. Ann. Myc. 15 (1917) 261.
On Citrus maxima, Reinking, Philip. Journ. Sci. 13 (1918) 165.
On coccids. Reinking, Phytopath. 9 (1919) 119.
LEPTOSTROMATACE^E
DIEDICKEA SINGULARIS Syd.
On Polyosma philippinensis. Ann. Myc. 15 (1917) 260.
On Polyosma sorsogonensis. Leafl. Philip. Bot. 9 (1925) 3137.
LASIOTHYRIUM CYCLOSCHIZON Syd.
On Aegiceras corniculatum. Philip. Journ. Sci. 12 (1917).
LEPTOTHYRIUM CIRCUMSCISSUM Syd.
On Mangifera indica. Baker, Philip. Agr. & For. 3 (1914) 162;
Reinking, Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9 (1919)
127.
MELANCONIALES
MELANCONIACE^E
COLLETOTRICHUM ARECAE Sydow.
On Areca catechu. Baker, Philip. Agr. & For. 4 (1914) 158; Rein-
king, Philip. Journ. Sci. 13 (1918) 165.
528 The Philippine Journal of Science 1931
COLLETOTKICHUM AKECAE Syd. Forma setis perpaucis praedita.
On Areca catechu. Ann. Myc. 15 (1917) 262.
COLLETOTKICHUM EUCHROUM Syd.
On Euphorbia neriifolia. Baker, Philip. Agr. & For. 3 (1914) 161.
COLLETOTRICHUM FALCATUM Went.
On Saccharwm officinarum. Phytopath. 9 (1919) 134; Philip. Agr.
Rev. 14 (1921) 431.
COLLETOTKICHUM GLOEOSPORIOIDES Penz.
On Citrus maxima. Philip. Journ. Sci. 13 (1918) 166; Philip. Agr.
9 (1920-21) 139; Philip. Agr. Rev. 14 (1921) 424.
COLLETOTKICHUM LUSSONIENSE Sacc.
On Manihot utilissima. Baker, Philip. Agr. & For. 3 (1914) 162;
Reinking, Philip. Journ. Sci. 13 (1918) 168; Phytopath. 9 (1919)
128.
COLLETOTKICHUM NIGRUM Ellis and Halsted.
On Capsicum annuum. Phytopath. 9 (1919) 117; Philip. Agr. 13
(1924-25) 165; 14 (1925-26) 500.
COLLETOTRICHUM PAPAYAE (Henn.) Syd.
On Carica papaya. Baker, Philip. Agr. & For. 3 (1914) 159; Ann.
Myc. 15 (1917) 262; Reinking, Philip. Journ. Sci. 13 (1918) 166;
Phytopath. 9 (1919) 118; Ann. Myc. 21 (1923) 105.
GLOEOSPORIUM MACROPHOMOIDES Sacc.
On Sesamum indicum. Baker, Philip. Agr. & For. 3 (1914) 164.
GLOEOSPORIUM AFFINE Sacc.
Saccardo, Syll. Fung. 3: 709.
On Hoya. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 29 — Los Ba-
nos (Baker $895).
GLOEOSPORIUM ALCHORNEAE Syd.
On Alchomea javanica. Ann. Myc. 15 (1917) 261.
On Alchomea rugosa. Leafl. Philip. Bot. 9 (1925) 3138.
GLOEOSPORIUM ALSTONIAE Sacc.
On Alstonia scholaris. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
29— Los Baiios (Baker 87S9).
GLOEOSPORIUM CANAVALIAE Syd.
On Canavalia, Baker, Philip. Agr. & For. 3 (1914) 159.
On Canavalia gladiata. Reinking, Philip. Journ. Sci. 13 (1918) 166.
On Phaseolus spp. Reinking, Phytopath. 9 (1919) 132.
GLOEOSPORIUM CATECHU Syd.
On Areca catechu. Baker, Philip. Agr. & For. 3 (1914) 158; Rein-
king, Philip. Journ. Sci. 13 (1918) 165.
GLOEOSPORIUM LEBBEK Syd.
On Albizzia lebbek. Ann. Myc. 15 (1917) 261.
46'3 Baker: Loiver Fungi 529
GLOEOSPORIUM MACROPHOMOIDES Sacc.
On Sesamum orientale (Sesamum indicum). Baker, Philip. Agr. &
For. 4 (1914) 164; Reinking, Philip. Journ. Sci. 13 (1918) 169;
Phytopath. 9 (1919) 136.
On Dioscorea esculenta. Reinking, Philip. Journ. Sci. 13 (1918)
166.
GLOEOSPORIUM MUSARUM Cke. and Mass.
On Musa sapientum* Philip. Agr. 10 (1922) 419; Philip. Agr. Rev.
18 (1925) 581; Philip. Agr. 13 (1924-25) 340.
GLOEOSPORIUM PALMARUM Oud.
On Areca catechu. Reinking, Philip. Journ. Sci. 13 (1918) 165.
GLOEOSPORIUM VANILLAE Cke.
On OrchidacesB. Baker, Philip. Agr. & For. 3 (1914) 163.
On Vanilla sp. Ann. Myc. 15 (1917) 261.
MARSONIA PAVONINA Syd.
On Macaranga sp. Ann. Myc. 15 (1917) 262.
On Macaranga utilis. Leafl. Philip. Bot. 9 (1925) 3138.
MELANCONIUM SACCHARI Cooke.
Saccardo, Syll. Fung. 14: 1019.
On Saccharum offlcinarum. Saccardo, Nuovo Giorn. Bot. Ital. 23
(1916) 29 — Ube, Mount Banahao (Baker 4-293, err. 3867); Baker,
Philip. Agr. & For. 5 (1916) 76, 343; Philip. Agr. Rev. 11 (1918)'
276; Reinking, Philip. Journ. Sci. 13 (1918) 169; Phytopath. 9
(1919) 134; Philip. Agr. Rev. 14 (1921) 429.
On Saccharum spontaneum. Ann. Myc. 15 (1917) 262.
PESTALOZZIA FUNEREA Desm.
On Carissa arduina. Baker, Philip. Agr. & For. 3 (1914) 162; 5
(1916) 74 — Los Bafios; Saccardo, Syll. Fung. 3 (1916) 791;
Nuovo Giorn. Bot. Ital. 23 (1916) 29— Los Banos (Baker 3788,
3 89 4).
PESTALOZZIA PALMARUM Cke. and Grev.
On Areca catechu. Baker, Philip. Agr. & For. 3 (1914) 160; 5
(1916) 73— Los Banos; Saccardo, Syll. Fung. 3 (1916) 796; Nuovo
Giorn. Bot. Ital. 23 (1916) 30— Los Banos (Baker 381J,).
On Cocos nucifera. Ann. Myc. 15 (1917) 262; Reinking, Phytopath.
9 (1919) 121; Philip. Agr. Rev. 14 (1921) 428; 18 (1925) 591.
PESTALOZZIA PAUCISETA Sacc.
On Uvaria. Ann. Myc. 15 (1917) 262.
On Mangifera indica. Reinking, Philip. Journ. Sci. 13 (1918) 167.
SEPTOGLOEUM ARACHIDIS Rac.
On Arachis hypogaea. Reinking, Journ. Sci. 13 (1918) 165; Phyto-
path. 9 (1919) 116.
263774 15
530 The Philippine Journal of Science «3i
HYPHALES
MUCEDINACEiE
ASPERGILLUS DELACRIOIXI Sacc. and Syd.
On Theobroma cacao. Baker, Philip. Agr. & For. 3 (1914) 164;
4 (1915) 165; Reinking., Philip. Journ. Sci. 13 (1918) 169; Phyto-
path. 9 (1919) 138.
ASPERGILLUS FLAVUS Link.
On fiber of Musa textilis. Philip. Journ. Sci. 32 (1927) 79.
ASPERGILLUS PERICONIOIDES Sacc.
On Carica papaya. Baker, Philip. Agr. & For. 3 (1914) 159; Rein-
king, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919) 118.
MYCOGNE CERVINA Ditm. var. THEOBROMAE Sacc.
On Theobroma cacao. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
30— Los Bafios (Baker 3884) ; Baker, Philip. Agr. & For. 5 (1916)
77; Reinking, Philip. Journ. Sci. 13 (1918) 169; Phytopath. 9
(1919) 138.
OIDIUM ERYSIPHOIDES Fr.
On Heliotropus indicus. Ann. Myc. 15 (1917) 263.
OOSPORA CANDIDULA Sacc.
Saccardo, Syll. Fung. 4: 12.
On Theobroma cacao. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
30 — Los Bafios (Baker 3729b) ; Baker, Philip. Agr. & For. 5
(1916) 77; Reinking, Philip. Journ. Sci. 13 (1918) 245; Phytopath.
9 (1919) 138.
OOSPORA HYALINULA Sacc. var. SORDIDULA Sacc.
On Capparis horrida. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
30— Los Bafios (Baker 3787d, err. 3887).
OOSPORA ORYZETORUM Sacc.
On Oryza sativa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 30 —
Los Banos (Baker 3867); Baker, Philip. Agr. & For. 5 (1916) 75;
Reinking, Philip. Journ. Sci. 13 (1918) 228; Phytopath. 9 (1919)
131.
RAMULARIA CATAPPAE Rac.
On Terminalia catappa. Baker, Philip. Agr. & For. 3 (1914) 164;
Reinking, Phytopath. 9 (1919) 138.
DEMATIACEiE
CERCOSPORA ACEROSUM Dickh. and Hem.
On Saccharum officinarum* Baker, Philip. Agr. & For. 4 (1914)
164.
CERCOSPORA APII Fres.
On Apium graveolens. Ann. Myc. 15 (1917) 264; Philip. Journ. Sci.
13 (1918) 165; Philip. Agr. 10 (1922) 349.
46,3 Baker: Lower Fungi 531
CERCOSPORA ARMORACIAE Sacc.
On Brassica spp. Baker, Philip. Agr. & For. 3 (1914) 159.
On Brassica pekinensis. Reinking, Philip. Journ. Sci. 13 (1918)
165.
On Brassica chinensis. Reinking, Phytopath. 9 (1919) 117.
CERCOSPORA ARTOCARPI Syd.
On Artocarpus incisa. Baker, Philip. Agr. & For. 3 (1914) 158;
Reinking, Phytopath. 9 (1919) 116.
On Artocarpus communis. Reinking, Philip. Journ. Sci. 13 (1918)
178.
CERCOSPORA OVERRHOI Welles.
On Averrhoa carambola. Welles, Philip. Journ. Sci. 19 (1921) 749.
CERCOSPORA BAUHINIAE Syd.
On Bauhinia malabarica. Ann. Myc. 15 (1917) 264.
CERCOSPORA BETICOLA Sacc.
On Beta vulgaris. Phytopath. 9 (1919) 116; Philip. Agr. 10 (1922)
349.
CERCOSPORA BRASSICOLA Heim.
On Brassica sinensis. Hennings, Engl. Jahrb. 37 (1905) 166 — Ja-
pan; Saccardo and Trotter, Syll. Fung. 22 (1913) 1413; Ann. Myc.
15 (1917) 264; Reinking, Phytopath. 9 (1919) 117.
On Brassica pekinensis. Reinking, Philip. Journ. Sci. 13 (1918)
165.
CERCOSPORA CANAVALIAE Syd.
On Canavalia gladiala. Baker, Philip. Agr. & For. 3 (1914) 159;
Reinking, Philip. Journ. Sci. 13 (1918) 165.
On Phaseolus sp. Reinking, Phytopath. 9 (1919) 132.
CERCOSPORA COFFEICOLA Berk, and Cooke.
On Coffea spp. Welles, Philip. Journ. Sci. 19 (1921) 743.
CERCOSPORA CRUENTA Sacc.
On Phaseolus aureus. Welles, Phytopath. 14 (1924) 357.
CERCOSPORA DUDDIAE Welles.
On Allium sativum and A. cepa. Welles, Phytopath. 13 (1923) 364*
CERCOSPORA GLIRICIDIAE Syd.
On Gliricidia sepium. Ann. Myc. 15 (1917) 264; Philip. Journ. Sci.
12 (1917) 380; 13 (1918) 382.
CERCOSPORA HENNINGSII Allesch.
On Manihot utilissima. Baker, Philip. Agr. & For. 4 (1914) 162;
Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919)
128.
CERCOSPORA LACTUCAE Stevenson. (Cercospora lactucae Welles.)
On Lactuca sativa. Welles, Phytopath. 13 (1923) 2S9.
CERCOSPORA LITSEAE-GLUTINOSAE Syd.
On Litsea glutinosa. Ann. Myc. 15 (1917) 264.
532 The Philippine Journal of Science 1931
CERCOSPORA LUSSONIENSE Sacc.
On Phaseolus lunatus. Baker, Philip. Agr. & For. 3 (1914) 1G3.
On Phaseolus spp. Reinking, Philip. Journ. Sci. 13 (1918) 165;
Phytopath. 9 (1919) 132.
CERCOSPORA MANGIFERAE Koord.
On Mangifera indica. Baker, Philip. Agr. & For. 3 (1914) 162; Ann.
Myc. 15 (1917) 264; Reinking, Philip. Journ. Sci. 13 (1918) 165;
Phytopath. 9 (1919) 127.
CERCOSPORA MANIHOTIS P. Henn.
On Manihot utilissima. Ann. Myc. 15 (1917) 265; Reinking, Philip.
Journ. Sci. 13 (1918) 165; Ann. Myc. 21 (1923) 106.
CERCOSPORA MELONGENAE Welles.
On Solanum melongena. Welles, Phytopath. 12 (1922) 63.
CERCOSPORA NICOTIANAE Ell. and Evht.
On Nicotiana tabacum* Baker, Philip. Agr. & For. 3 (1914) 162;
Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919)
117; Ann. Myc. 21 (1923) 106; Philip. Agr. Rev. 18 (1925) 570;
Philip. Agr. 15 (1926) 300.
CERCOSPORA OCCIDENTALIS Cke. var. CASSIOCARPA Sacc.
On Cassia occidentale. Ann. Myc. 15 (1917) 265.
CERCOSPORA PACHYDERMA Syd.
On Dioscorea spp. Baker, Philip. Agr. & For. 4 (1914) 161; Rein-
king, Phytopath. 9 (1919) 124.
On Dioscorea alata. Ann. Myc. 15 (1917) 265.
On Dioscorea esculenta. Reinking, Philip. Journ. Sci. 13 (1918)
165.
CERCOSPORA PAHUDIAE Syd.
On Pahudia romboidea. Baker, Philip. Agr. & For. 3 (1914) 163.
CERCOSPORA PANTOLEUCA Syd.
On Clitoria ternatea. Baker, Philip. Agr. & For. 3 (1914) 160.
CERCOSPORA PERSONATA (B. and C.) Ell.
On Arachis hypogaea. Baker, Philip. Agr. & For. 3 (1914) 158; Ann.
Myc. 15 (1917) 265; Philip. Journ. Sci. 12 (1917) 380; Reinking,
Phytopath. 9 (1919) 115.
CERCOSPORA PUERARIAE Syd.
On Pueraria sp. Ann. Myc. 15 (1917) 265.
CERCOSPORA SESAMI A. Zimm.
On Sesamum orientate (S. indicum). Baker, Philip. Agr. & For. 3
(1914) 164; Philip. Agr. & For. 6 (1917) 294; Ann. Myc. 15 (1917)
265; Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9
(1919) 136.
CERCOSPORA STIZOLOBII Syd.
On Mucuna deeringiana (Stizolobium deeringianum) . Baker, Philip.
Agr. & For. 3 (1914) 164; Reinking, Philip. Journ. Sci. 13 (1918)
165.
On Stizolobium niveum. Reinking, Phytopath. 9 (1919) 132.
46,3 Baker: Lower Fungi 533
CERCOSPORA SUBSESSILIS Syd.
On Melia azedarach. Ann. Myc. 15 (1917) 265.
CERCOSPORA TIGLII Henn.
On Croton tiglium. Baker, Philip. Agr. & For. 4 (1914) 161.
CERCOSPORA UBI Rac.
On Dioscorea spp. Baker, Philip. Agr. & For. 3 (1914) 161; Reinking,
Phytopath. 9 (1919) 124.
On Dioscorea esculenta. Reinking, Philip. Journ. Sci. 13 (1918) 165.
ALTERNARIA BRASSICAE (Berk.) Sacc.
On Brassica culta. Ann. Myc. 15 (1917) 266.
CERCOSPORINA CARTHAMI Syd.
On Carthamus tinctorium. Baker, Philip. Agr. & For. 3 (1914) 159.
CLADOSPORIUM HERBARUM L.
On Phaseolus limatus. Baker, Philip. Agr. & For. 3 (1914) 163;
Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919)
132.
CLADOSPORIUM LINEOLATUM Sacc.
On Capparis micracantha. Ann. Myc. 15 (1917) 264.
CLASTEROSPORIUM MAYDICUM Sacc.
On Zea mays, Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 31 — Los
Bafios {Baker 8733a); Baker, Philip. Agr. & For. 5 (1916) 78;
Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919)
140.
CONIOSPORIUM BAMBUSAE (Thuem. and Bolle) Sacc.
On Bambusa sp. Ann. Myc. 15 (1917) 263.
On Bambusa longinodis. Ann. Myc. 21 (1923) 105.
CONIOSPORIUM EXTREMORUM Syd.
On Saccharum officinarum. Baker, Philip. Agr. & For. 3 (1914)
164; 5 (1916) 343; Philip. Agr. Rev. 11 (1918) 276; Reinking,
Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919) 136.
CONIOSPORIUM ORYZINUM Sacc.
On Oryza sativa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 31 —
Los Baiios (Baker 8773); Baker, Philip. Agr. & For. 5 (1916) 76;
Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919)
131.
CONIOSPORIUM UNILATERALE Sacc. and Peyr.
On Schizo8tachyum sp. Ann. Myc. 15 (1917) 263.
CONIOSPORIUM VINOSUM (B. and C.) Sacc.
On Saccharum officinarum. Baker, Philip. Agr. & For. 3 (1914) 164;
5 (1916) 343; Reinking, Philip. Journ. Sci. 13 (1918) 165; Philip.
\ Agr. Rev. 11 (1918) 276; Phytopath. 9 (1919) 136.
534 The Philippine Journal of Science 1931
DICHOTOMELLA AREOLATA Sacc.
On Artocarpus Integra (A. integrifolia) . Baker, Philip. Agr. & For.
3 (1914) 158; Reinking, Philip. Journ. Sci. 13 (1918) 165; Phyto-
path. 9 (1916) 116.
HELMINTHOSPORIUM CARYOPSIDUM Sacc.
On Andropogon sorghum (Sorghum vulgare, Holcus sorghum) . Baker.
Philip. Agr. & For. 3 (1914) 164; Saccardo, Nuovo Giorn. Bot. Ital.
23 (1916) 32— Los Bafios (Baker 875k, $808, 3812) ; Baker, Philip.
Agr. & For. 5 (1916) 77; Reinking, Philip. Journ. Sci. 13 (1918)
165; Phytopath. 9 (1919) 137.
HELMINTHOSPORIUM CURVULUM Sacc.
On Zea mays. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 32 — Los
Bafios (Baker 3733b); Baker, Philip. Agr. & For. 5 (1916) 78; Re-
inking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919) 140.
HELMINTHOSPORIUM FICINUM Sacc. (Helminthosporium ficinum Yates.)
On Ficus caudatifolia. Philip. Journ. Sci. 13 (1918) 382; Ann. Myc.
20 (1922) 73.
On Ficus. Ann. Myc. 21 (1923) 105; Leafl. Philip. Bot. 9 (1925)
3138.
HELMINTHOSPORIUM INCONSPICUUM C. and Ell.
On Zea mays. Philip. Agr. Rev. 4 (1911) 357; Baker, Philip. Agr. &
For. 3 (1914) 164; Ann. Myc. 15 (1917) 265; Reinking; Philip.
Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919) 140; Philip. Agr.
12 (1923-24) 457; Philip. Agr. Rev. 18 (1925) 571; Philip. Agr.
15 (192'5) 127.
HELMINTHOSPORIUM INVERSUM Sacc.
On Erythrina indica. Ann. Myc. 15 (1917) 265.
HELMINTHOSPORIUM ORYZAE Breda de Haan.
On Oryza sativa. Ocfemia, Phytopath. 12 (1922) 34; Am. Journ.
Bot. 11 (1924) 437.
HELMINTHOSPORIUM PAPAYAE Syd.
On Carica papaya. Sydow, Ann. Myc. 21 (1923) 105.
HELMINTHOSPORIUM RAVENELII Berk, and Curt.
On Sporobolus elongatus. Ann. Myc. 15 (1917) 266; 21 (1923) 105.
On Panicum auritum. Philip. Journ. Sci. 13 (1918) 383.
On Sporobolus sp. Leafl. Philip. Bot. 9 (1925) 3138.
HADRONEMA ORBICULARE Sydow.
On Quercus sp. Philip. Journ. Sci. 12 (1917) 380; 13 (1918) 382.
PERICONIA PHILIPPINENSIS Sacc.
On Panicum. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 32— Los
Banos (Baker 3766).
SARCINELLA RAIMUNDOI Sacc.
On Solanum melongena. Baker, Philip. Agr. & For. 3 (1914) 164;
Reinking, Philip. Journ. Sci. 13 (1918) 165; Phytopath. 9 (1919)
136.
46, 3 Baker: Lower Fungi 535
SEPTONEMA PHILIPPINUM Sacc.
On Imperata cylindrica. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
32— Los Banos (Baker 3769).
SPORODESMIUM BAKERI Syd.
On Musa sapientum. Baker, Philip. Agr. & For. 3 (1914) 162;
Reinkinig, Philip. Journ. Sci. 13 (1918) 165.
On Musa paradisiaca sapientum. Reinking, Phytopath. 9 (1919) 129.
TORULA DICHROA Sacc.
On Saccharum spontaneum. Saccardo, Nuovo Giorn. Bot. Ital. 23
(1916) 31— Los Banos (Baker 3737).
TORULA HERBARUM Link.
Saccardo, Syll. Fung. 4: 256.
On Capparis horrida. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
30— Los Banos (Baker 3787c).
TORULA HERBARUM Lk. f. QUATERNELLA Sacc.
On Thunbergia grandiflora. Ann. Myc. 15 (1917) 263.
TRICHOSPORIUM COCCIDICOLA Sacc.
On Phenacuspis mischocarpi and Mischocarpus fuscesce?is. SACCARDO,
Nuovo Giorn. Bot. Ital. 23 (1916) 31— Mount Maquiling (Baker
3859).
STIGMELLA MANILENSIS Sacc.
On Allophyllum dimorphum. Ann. Myc. 15 (1917) 268.
ZYGOSPORIUM OSCHEOIDES Mont.
On Areca catechu. Reinking, Philip. Journ. Sci. 13 (1918) 165.
TUBERCULARIACE^
DENDRODOCHIUM LUSSONENSE Sacc.
Ann. Myc. 15 (1917) 267.
EXOSPORIUM DURUM Sacc.
On Cocos nucifera. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 33 —
Ube, Mount Banahao (Baker 386b) ; Baker, Philip. Agr. & For.
5 (1916) 74 — Mount Banahao; Reinking, Philip. Journ. Sci. 13
(1918) 165; Reinking, Phytopath. 9 (1919) 121; Philip. Agr. Rev.
18 (1925) 591.
EXOSPORIUM PULCHELLUM Sacc.
On Areca catechu. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 33 —
Los Banos (Baker 3753, 3799) ; Baker, Philip. Agr. & For. 5 (1916)
73 — Los Banos; Reinking, Philip. Journ. Sci. 13 (1918) 165.
On Omnia palindan. Ann. Myc. 15 (1917) 266.
FUSARIUM CUBENSE Efs.
On Musa sapientum. Philip. Agr. Rev. 13 (1920) 128; Phytopath. 10
(1920) 504.
On Musa textilis. Philip. Agr. Rev. 16 (1923) 106; Lee and Serrano,
Phytopath. 13 (1923) 354; Philip. Agr. 19 (1930) 27.
536 The Philippine Journal of Science
FUSARIUM THEOBROMAE App. and Strunk.
Saccardo, Syll. Fung. 18: 672.
On Theobroma cacao. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916)
33 — Los Banos (Baker 3885); Reinking, Philip. Journ. Sci. 13
(1918) 165; Phytopath. 9 (1919) 138.
HYMENOPSIS CUDRANIAE Mass.
On Cudrania javanica. Philip. Journ. Sci. 13 (1918) 384.
HYMENULA COPELANDI Sacc.
On Diospyrus sp. Ann. Myc. 15 (1917) 267.
ILLOSPORIUM TABACINUM Sacc.
On Macaranga. Saccardo, Ann. Myc. 13 (1915) 128 — Los Banos
(Baker 33 22).
PIONNOTES CAPILLACEA Sacc.
On Persea americana and P. gratissima. Saccardo, Nuovo Giorn. Bot.
Ital. 23 (1916) 34— Los Banos (Baker 3816); Baker, Philip. Agr.
& For. 5 (1916) 76.
SPEGAZZINIA MELIOLAE A. Zimm.
On Meliola callicarpae. Philip. Journ. Sci. 12 (1916) 363; Ann. Myc.
15 (1917) 268.
SPEGAZZINIA ORNATA Sacc.
Saccardo, Syll. Fung. 4 (1917) 758.
On Oryza sativa. Saccardo, Nuovo Giorn. Bot. Ital. 23 (1916) 32 —
Los Banos (Baker 3772, 3770 , 3803) ; Baker, Philip. Agr. & For.
5 (1916) 76— Los Banos; Reinking, Philip. Journ. Sci. 13 (1918)
165; Phytopath. 9 (1919) 131.
GRAPHIOLA ARENGAE Rac.
On Arenga ambong. Ann. Myc. 15 (1917) 178.
GRAPHIOLA CYLINDROSPORA Syd.
On Livistonia. Philip. Agr. & For. 5 (1916) 74.
MYCELIA STERILIA
OZONIUM GLUMICOLA Sacc.
On Schizostachum acutiflorum. Saccardo, Nuovo Giorn. Bot. Ital.
23 (1916) 34— Mount Maquiling (Baker 3813).
SCLEROTIUM ROLFSII Sacc.
On Nicotiana tabacum. Philip. Agr. Rev. 14 (1921) 427; Philip. Agr.
15 (1926) 290.
On Ly coper sicwm esculentum and Capsicum annuum. Philip. Agr. 13
(1924-25) 166; 15 (1926) 580.
On seedlings. Philip. Agr. Rev. 18 (1925) 564.
On Oryza sativa. Philip. Agr. 15 (1926) 362; Philip. Agr. Rev. 19
(1926) 238.
The Philippine
Journal of Science
Vol. 46 DECEMBER, 1931 No. 4
WORM PARASITES OF THE BROWN RAT (MUS NORVE-
GICUS) IN THE PHILIPPINE ISLANDS, WITH
SPECIAL REFERENCE TO THOSE FORMS
THAT MAY BE TRANSMITTED TO
HUMAN BEINGS
By Marcos A. Tubangui
Of the Division of Biology and Serum Laboratory
Bureau of Science, Manila
NINETEEN TEXT FIGURES
INTRODUCTION
The role of rats as carriers and reservoirs of bubonic plague
and other bacterial as well as spirochetal infections, has long
been well recognized. For this reason various antirat meas-
ures have been in vogue in different parts of the world, espe-
cially in seaports, for the control and prevention of these
diseases. The fact, however, that these animals are often in-
fested with certain parasitic worms that are also a menace to
human health, is not so well known. For this reason and be-
cause of the fact that the helminthic fauna of rats in the Philip-
pine Islands has never been studied to any great extent, it seemed
worth while to undertake a systematic examination of these
animals in order to determine their parasites and to find if they
harbor forms that are transmissible to man.
BATS EXAMINED AND THE INCIDENCE OF INFESTATION
The survey was limited to the brown or Norway rat, Mus nor-
vegicus Erxleben, 1777 (= M. decumanus Pallas, 1778), since
this was the only rat constantly available in large numbers.
A total of nine hundred fifty of these rodents were dissected
264209 ggfT
538 The Philippine Journal of Science 1931
during the period from May 7, 1930, to January 14, 1931. They
were trapped in the different sections of the City of Manila
and were among those sent to the Bureau of Science by the
Philippine Health Service for routine bubonic-plague inspection.
A list of the different parasites encountered and their incidence
are given in Table 1. One species of roundworm, Syphacia
obvelata, is not represented in the table, but it is believed to
infest rats in the Philippines in view of its having been reported
by Riley (1919) in a child residing in Zamboanga, Mindanao.
With the exception of the flukes, a new species of the cestode
genus Raillietina, and a new nematode in the genus Rictularia,
all of which are apparently restricted to the Philippines in their
distribution, the different worms collected have been reported
from other countries. The following were the most commonly
met with in the order they are named: The larval form of
Tsenia tseniaformis (commonly known as Cysticercus fasciola-
ris), Hepaticola hepatica, Raillietina garrisoni sp. nov., Strongy-
loides ratti, Hymenolepis diminuta, Nippostrongylus muris,
Trichosomoides crassicauda, and Gongylonema neoplasticum.
Hymenolepis nana and Heterakis spumosa, which are common
in rats in many countries, were rarely encountered. Trichi-
nella spiralis, the most dangerous worm of rats from the pub-
lic-health standpoint, was not found at all.
Table 1. — Parasites encountered in nine hundred fifty rats.
Name of parasites. Infestation.
Per cent.
Trematodes :
Euparyphium ilocanum 0.5
Euparyphium guerreroi 0.1
Euparyphium murmwm sp. nov. 0.1
Cestodes :
Taenia tseniaformis (larval form) 94.0
Raillietina garrisoni sp. nov. 86.0
Hymenolepis diminuta 64.0
Hymenolepis nana 1.7
Nematodes :
Gongylonema neoplasticum 44.0
Hepaticola hepatica 90.0
Heterakis spumosa 0.4
Nippostrongylus muris 58.0
Protospirura mwricola 1.3
Rictularia whartoni sp. nov. 0.4
Strongyloides ratti 74.0
Trichosomoides crassicauda 57.0
Acanthocephala :
Moniliformis moniliformis 4.2
46,4 Tubangui: Worm Parasites of the Brown Rat 539
The incidence of the worms did not seem to depend upon the
time of the year but rather, in the case of the flukes, at least,
on the environment of their hosts. It was noticed at the termi-
nation of the survey that these particular parasites were ob-
tained only from some of the rats that were trapped inside the
piers of Manila Bay and in the immediate neighborhood of the
landing places of boats along Pasig River. This may be regard-
ed as purely accidental, but it may also mean that either the in-
termediate hosts of these flukes, which most probably are snails,
exist in some of the bodies of water in Manila or the rats that
harbored them might have been brought to the city from other
localities on board of ships and boats. The matter deserves
further inquiry.
DESCRIPTIONS OF PARASITES
The parasites determined represent two phyla in the animal
kingdom, namely, the Platyhelminthes, or flatworms, and the
Nemathelminthes, or roundworms. The flukes (class Trema-
toda) and the tapeworms (class Cestoda) are members of the
phylum Platyhelminthes, while the so-called true roundworms
(class Nematoda) and the proboscis worm (class Acanthoce-
phala) belong to the Nemathelminthes.
Phylum PLATYHELMINTHES Claus, 1885
Class TREMATODA Rudolphi, 1808
Subclass DIGENEA v. Beneden, 1858
Order PROSOSTOMATA Odhner, 1905
Suborder DISTOMATA Zeder, 1800
Superfamily ECHINOSTOMATOIDEA Faust, 1929
Family ECHINOSTOMATIDiE Looss, 1902
Subfamily ECHINOSTOMATIN^E Looss, 1899
Genus EUPARYPHIUM Dietz, 1909
EUPARYPHIUM ILOCANUM (Garrison, 1908) Tubangui, 1931. fi*. 1.
Synonyms: Fascioletta ilocana Garrison, 1908; Echinostoma Mocanwm
(Garrison, 1908) Odhner, 1911.
For many years this fluke was regarded as a parasite peculiar
to man in the northwestern provinces of Luzon, Philippine Is-
lands. Its occurrence in rats has been only recently demon-
strated by the present writer (Tubangui, 1931). In the survey
540
The Philippine Journal of Science
1931
FiG. 1. Euparyphium Uocanum.
a, Entire worm, ventral view; 6,
anterior end, showing arrange-
ment of spines on cephalic collar,
ventral view. (After Tuhangui,
1931.)
on which this report is based, five
or a little more than 0.5 per cent of
the nine hundred fifty rats examined
were infested with it.
Description. — B o d y moderately
large, elongate, 5.57 to 8.02 millime-
ters in length by 1.33 to 1.58 milli-
meters in maximum breadth at or
near the equator of body. Lateral
sides of body from anterior end to
acetabulum rolled ventrally. Cuticle
armed with flat scalelike structures
distributed ventrally from anterior
end to second testis or slightly be-
yond that level, and dorsally from
anterior end to anterior level of ace-
tabulum; scales 13.5 to 24.7 by 13.5
to 18.0 microns in size, those at an-
terior end being smaller. Suckers
close together ; oral sucker small, sub-
terminal, 0.19 to 0.24 millimeter in
transverse diameter ; acetabulum
large, cup-shaped, at middle of an-
terior third of body length, 0.60 to
0.69 by 0.64 to 0.74 millimeter in size.
Oral sucker surrounded dorsally and
laterally by a collar (fig. 1, 6) bear-
ing fifty-one spines arranged in two
alternating rows; collar 0.38 to 0.46
millimeter in diameter, reniform, its
two rounded ventral angles united by
a narrow ridge. Collar spines may
be grouped as follows: Six ventral
corner spines on each side of collar,
the smallest of which measures 36.0
by 11.2 microns, the broadest 42.7 by
15.7 microns, and the longest 45.0
by 11.2 microns; fourteen lateral
spines on each side, arranged in pairs
and eleven dorsal spines; lateral and
dorsal spines 31.5 to 45.0 by 11.2 to
13.5 microns in size.
46,4 Tubangui: Worm Parasites of the Brown Rat 541
Mouth terminal to subterminal, followed occasionally by pre-
pharynx 0.03 to 0.05 millimeter in length; pharynx 0.19 to 0.20
by 0.15 to 0.17 millimeter in size; oesophagus 0.10 to 0.20 milli-
meter long, bifurcating immediately in front of genital pore,
midway between pharynx and acetabulum or slightly anterior
of that level; intestinal caeca reach posteriorly to from 0.24 to
0.43 millimeter from posterior end of body.
Testes tandem, postequatorial, at third fourth of body length,
either elongate and each divided into anterior and posterior
lobes by transverse constriction or shorter and distinctly 3- to
4-lobed. Cirrus sac large, 0.51 to 0.65 by 0.26 to 0.34 millimeter
in size, reaching to but not extending posteriorly beyond equa-
tor of acetabulum; incloses prominent seminal vesicle, well-de-
veloped pars prostatica, and long protrusible cirrus. Common
genital opening preacetabular, behind oesophageal bifurcation,
to one side of median line.
Ovary globular or slightly compressed transversely, median,
pretesticular, usually behind middle of second fourth of body
length, 0.31 to 0.43 by 0.34 to 0.48 millimeter in size; shell gland
between ovary and anterior testis ; receptaculum seminis absent,
Laurer's canal present; uterus well developed, occupying space
bounded by ovary, acetabulum, and intestinal caeca. Vitellaria in
moderately large follicles, commencing anteriorly on both sides
at level about midway between posterior border of acetabulum
and anterior border of ovary; anteriorly they are extracaecal,
but behind second testis the follicles from the two sides unite
and occupy most of posterior region of body; transverse vitel-
line ducts and vitelline reservoir dorsal of shell gland, directly
in front of anterior testis. Eggs, numerous, operculated, light
brown or yellowish, 85.5 to 101.5 by 54.0 to 65.2 microns in size.
Excretory system typical of echinostomes in general; excre-
tory bladder long, with several small side branches, dividing
into two principal branches behind second testis ; excretory pore
at extreme posterior end of body.
Location. — Small intestine.
Life history. — Unknown. It is most probable, however, from
what is known of the life history of mammalian trematodes
that the intermediate host is a fresh-water snail. It might be
interesting to note moreover that the eercariae of related flukes
assume the infective stage by encysting within their own rediae
or in the tissues of their intermediate hosts; others encyst on
plants, fishes, or in tadpoles.
542
The Philippine Journal of Science
1931
Fig. 2. Euparyphium guerreroi.
a, Entire worm, ventral view; b,
anterior end, showing arrange-
ment of spines on cephalic collar,
ventral view. (After Tubangui,
1931.)
Prevention. — Bearing in mind the
possible modes of infestation with
this parasite as noted in the discus-
sion of its life history, prevention
should consist in the avoidance of raw
or improperly cooked vegetables,
snails, and fishes and unboiled or un-
filtered surface water as food and
drink, respectively, especially in those
places where the fluke is known to
occur.
References.— 14, 16, 21, 31, 41, 51,
55.1
EUPARYPHIUM GUERREROI Tubangui, 1931.
Fig. 2.
Description. — Body slender, elon-
gate, measuring 2.92 to 4.03 milli-
meters in length by 0.37 to 0.50
millimeter in maximum breadth
across acetabulum or anywhere be-
tween this organ and anterior testis.
Cuticle armed with flat scales, dor-
sally from anterior end to level of
acetabulum and ventrally from ante-
rior end to posterior testis or slightly
beyond that level; scales 6.0 to 15.0
by 5.5 to 9.4 microns in size, anterior
ones being smaller. Oral sucker
small, subterminal, 0.10 to 0.12 milli-
meter in transverse diameter; aceta-
bulum larger, at middle of anterior
third of body length, 0.27 to 0.36 by
0.31 to 0.34 millimeter in size. Oral
sucker surrounded dorsally and lat-
erally by a collar (fig. 2, 6) bearing
fifty-five spines arranged in two al-
ternating rows; collar 0.22 to 0.26
millimeter across, reniform, its two
ventral angles united by a narrow
ridge. Collar spines may be grouped
xThe numbers refer to the list of refer-
ences, which are arranged alphabetically
and numbered, at the end of this paper.
46,4 Tubangui: Worm Parasites of the Brown Rat 543
as follows : Five ventral corner spines on each side of cephalic
collar, 24.7 to 31.5 by 9.0 to 11.9 microns; fifteen lateral spines
on each side, 27.0 to 29.2 by 9.0 microns; and fifteen dorsal
spines, 11.2 to 13.5 by 6.7 to 9.0 microns in size.
Mouth subterminal to terminal, followed by prepharynx 0.03
to 0.07 millimeter long; pharynx 0.10 to 0.11 by 0.07 to 0.08
millimeter in size; oesophagus 0.08 to 0.15 millimeter long, bi-
furcating in front of level of genital pore ; intestinal caeca long,
narrow in diameter, reaching from 0.21 to 0.24 millimeter from
posterior end of body.
Testes tandem, postequatorial, at third fourth of body length,
oval or sausage-shaped, often transversely constricted into ante-
rior and posterior lobes ; anterior testis usually smaller, at least
shorter, 0.19 to 0.36 by 0.15 to 0.22 millimeter in size; posterior
testis 0.27 to 0.39 by 0.12 to 0.20 millimeter in size. Cirrus
pouch oval, 0.17 to 0.27 by 0.10 to 0.13 millimeter in size, not
reaching posteriorly beyond equator of acetabulum; incloses
large seminal vesicle, moderately developed pars prostatica, and
protrusible cirrus. Common genital opening preacetabular, be-
hind oesophageal bifurcation, to one side of median line.
Ovary globular or slightly compressed, 0.10 to 0.15 by 0.07 to
0.13 millimeter in size, immediately preequatorial, pretesticu-
lar; shell gland prominent, filling most of the space between
ovary and anterior testis ; receptaculum seminis absent, Laurer's
canal present; uterus short, with few coils. Vitellaria in small
to moderately large follicles, commencing anteriorly at middle
of second fourth of body length, those on left side usually
commencing at a more posterior level; behind second testis
follicles from two sides unite and extend to posterior end of
body; transverse vitelline ducts and vitelline reservoir dorsal
of shell gland and immediately in front of first testis. Eggs
few, operculated, thin shelled, light brown or yellowish, 78.7
to 85.5 by 54.0 to 60.7 microns in size.
Excretory system of usual echinostome type; excretory blad-
der long, tubular, dividing into two branches behind second tes-
tis ; excretory pore at extreme posterior end of body.
Location. — Small intestine.
Life history. — Unknown.
Reference. — 55.
EUPARYPHIUM MURINUM sp. nov. Figf. 3.
The description of this parasite is based on the examination
of two lots of material. One lot, consisting of a small number
of specimens, is part of our collection and was obtained from a
544
The Philippine Journal of Science
1931
&m
wmi
wmm
rat that was at the same time infested
with E. ilocanum. The other lot con-
sisting of numerous specimens and
labelled "parasitos encontrados en el
intestino de un raton, Manila, Agosto,
1909," was collected by Dr. Luis Guer-
rero. It was kindly turned over to me
for determination by Dr. Onofre Gar-
cia who found it among the parasitolog-
ical collections of the University of
Santo Tomas, Manila. I take this op-
portunity to express my thanks to Doc-
tor Guerrero and Doctor Garcia.
This fluke differs from the two pre-
ceding species in the number of its
collar spines, of which there are forty-
five to forty-six, and in the position of
its cirrus pouch that extends posteriorly
beyond the equator of the acetabulum.
In the number of its collar spines it is
similar to Echinostoma gotoi Ando and
Ozaki, 1923, another rat trematode, but
again it may be distinguished from the
latter by the position of its cirrus sac
and also by the character of its uterus,
which is short and contains only a few
coils and eggs.
Description. — Body small, elongate,
2.65 to 4.50 by 0.45 to 0.65 millimeters
in size. Cuticle armed with flat scales,
dorsally from anterior end to acetabu-
lum and ventrally from anterior end
to posterior level of first testis or
slightly beyond. Oral sucker small,
subterminal, 0.10 millimeter in trans-
verse diameter ; acetabulum 0.32 to 0.42
by 0.23 to 0.32 millimeter in size, at
anterior fourth of body length. Head
collar reniform, 0.23 to 0.27 millimeter
across, bearing forty-five spines ar-
ranged in two alternating rows and measuring 37.5 to 44.2 by
a
Fig. 3. Euparyphium murinum
sp. nov. a, Entire worm,
ventral view ; b, anterior end,
showing arrangement of spines
on cephalic collar, ventral
view.
46,4 Tubangui: Worm Parasites of the Brown Rat 545
8.0 to 9.2 microns. Occasionally there are forty-six collar spines
due to the presence of a small accessory dorsal spine (fig. 3, b).
Mouth subterminal; prepharynx absent or very short; pha-
rynx oval, 0.10 to 0.13 by 0.07 to 0.09 millimeter in size; oeso-
phagus 0.07 to 0.12 millimeter long, bifurcating immediately
in front of level of genital pore; intestinal cseca long, reaching
to near posterior end of body.
Testes tandem, postequatorial, oval to sausage-shaped, with
smooth borders or slightly constricted at middle; anterior tes-
tis usually smaller, 0.32 to 0.48 by 0.16 to 0.25 millimeter in size ;
posterior testis 0.33 to 0.53 by 0.15 to 0.26 millimeter. Cirrus
pouch oval, 0.25 to 0.36 by 0.10 to 0.13 millimeter in size, usually
to one side of median line, dorsal to acetabulum and extending
posteriorly beyond the equator of this organ; incloses seminal
vesicle, pars prostatica, and protrusible cirrus. Common genital
pore immediately preacetabular, a little to one side of median
line.
Ovary globular or slightly transversely oval, preequatorial,
pretesticular, 0.10 to 0.15 millimeter in transverse diameter.
Shell gland conspicuous, between ovary and first testis. Recep-
taculum seminis absent, the distal portion of oviduct being di-
lated and probably functioning as seminal receptacle; Laurer's
canal present. Uterus short, with few coils. Vitelline glands
in the form of distinct follicles extending from 0.10 to 0.60
millimeter behind acetabular level to near posterior end of body.
Eggs few, oval, operculated, thin shelled, yellowish, 88.4 to 95.2
by 57.8 to 61.2 microns in size.
Excretory system of the usual echinostome type; excretory
bladder tubular, bifurcating behind second testis ; excretory pore
at extreme posterior end of body.
Specific diagnosis. — Euparyphium: Body elongate, 2.65 to 4.50
by 0.45 to 0.65 millimeters in size. Head collar 0.23 to 0.27
millimeter in transverse diameter, with forty-five spines
measuring 37.5 to 44.2 by 8.0 to 9.2 microns. Prepharynx
very short or absent, oesophagus 0.07 to 0.12 millimeter long.
Testes oval to sausage-shaped, with smooth borders or slightly
constricted at middle; cirrus sac oval, 0.25 to 0.36 by 0.10 to 0.13
millimeter in size, reaching posteriorly beyond equator of aceta-
bulum. Ovary globular or transversely oval, preequatorial;
vitellaria extend from 0.10 to 0.60 millimeter behind acetabular
level to posterior end of body. Eggs few, 88.4 to 95.2 by 57.8
to 61.2 microns in size.
546 The Philippine Journal of Science 1931
Location. — Small intestine.
Locality. — Manila, Philippine Islands.
Type specimens. — Philippine Bureau of Science parasitolo-
gical collection, No. 64; paratypes in parasitological collection
of the University of Santo Tomas, Manila.
Life history. — Unknown.
References.— 1, 10, 11, 13, 30, 31, 55.
Class CESTODA Rudolphi, 1808
Subclass CESTODA (s. str.) Monticelli, 1892
Order CYCLOPHYLLIDEA Braun,1900
Superfamily T7ENIOIDEA Zwicke, 1841
Family TiENIID JE Ludwig, 1886
Subfamily T^NIIN^ Stiles, 1896
Genus TiENIA Linnaeus, 1758
TAENIA T^NIAFORMIS (Batsch, 1786) Wolffhiisel, 1911. Fig. 4.
Synonym: Tsenia crassicollis Rudolphi, 1810.
The larval stage of this tapeworm is commonly known as
Cysticercus fasciolaris Eudolphi, 1808 (= Strobilocercus fascio-
laris Sambon, 1924). It is one of the commonest parasites
of the brown rat, the livers of 94 per cent of the animals ex-
amined being infested with it. The adult stage has so far
been found only in cats. Krabbe, according to Stiles (1906),
pointed out long ago that in Jutland sandwiches of chopped raw
mice were eaten by the common people for the relief of anuria
and suggested that this custom might be responsible for the
occasional presence of the parasite in man. Thus far, however,
no case of the sort has been reported.
Description. — The larvae are inclosed in globular cysts, partly
visible on the surface of the liver of infested rats as whitish
semitransparent areas. These cysts are 5 to 16 millimeters in
diameter and are easily separated from the hepatic tissue.
The larvae themselves are elongate, measuring 30 to 200 milli-
meters in length by 2 to 6 millimeters in maximum width near
the anterior end. The body (fig. 4, a) is strobilate, which char-
acter differentiates it from the other bladderworms {Cysti-
cercus species), for which reason Sambon (1924) proposed for
it the term Strobilocercus. In living specimens the anterior
portion is usually wider and thicker due to the contraction of
46,4 Tubangui: Worm Parasites of the Brown Rat 547
0.5 mm
h
Fig. 4. Tsenia t&niaformis. a,
Entire larva (after Sambon, 1924) ;
view ; c, rostellar hooks.
b, scolex, anterior
the body at this region and presents a terminal wedge-shaped
depression due to the invagination of the scolex. The posterior
end is usually more slender, terminating in a very much reduced
bladder. Sometimes the segmentation of the body is so distinct
and the length so great that this larvae has been mistaken for a
small mature tapeworm. Occasionally rudimentary reproduc-
tive organs are present among some of the segments. The
scolex (fig. 4, 6) is large, thick, 1.3 to 1.7 millimeters broad;
suckers prominent, cup-shaped, 0.32 to 0.38 millimeter in diam-
eter; rostellum short, columnar, 1.12 to 1.14 millimeter in diam-
eter, crowned with 26 to 52 hooks, according to various authors
(hooks of Philippine material 38 to 42). The hooks (fig. 4, c)
are of the characteristic shape found in the group of tapeworms
548 The Philippine Journal of Science 1931
to which this parasite belongs and are arranged in two concentric
circles; those forming the upper row are larger, 380 to 420
microns long, their free pointed ends being almost on a line
with those of the shorter hooks of the lower ring with which
they alternate; the smaller hooks are 250 to 270 microns long.
Location. — Liver.
Life history. — The encysted strobilocercus in the liver of rats
and other rodents represents the infective stage in the transmis-
sion of this parasite to its final host. If fed to a cat, the larva
is liberated in the small intestine, attaches itself to the intestinal
wall, increases in size, and, after two to three months, be-
comes mature. The eggs of the adult parasite escape with the
faeces of the host and, if these are ingested by a rat or any other
animal that can play the role of intermediate host, the inclosed
embryos are freed from their shells in the intestine. These
embryos on reaching the liver become encysted and are developed
into strobilocerci. They reach the liver presumably through the
circulatory system after penetrating through the intestinal wall.
References.— -20, 27, 37, 49.
Family DAVAINEIDJE Fuhrmann, 1907
Subfamily DAVAINEIN^E Braun, 1900
Genus RAILLIETINA Fuhrmann, 1920
RAILLIETINA GARRISONI sp. nov. Fig. 5.
Synonym: ? Davainea madagascariensis (Davaine) of Garrison, 1911.
This appears to be the commonest intestinal cestode infest-
ing the brown rat in the Philippines. It bears a close resem-
blance to R. celebensis, but differs from the latter, as described
by Janicki (1902) and by Meggitt and Subramanian (1927), in
having a larger number of testes and uterine egg capsules and
in the larger size of its cirrus pouch. It is, therefore, proposed
as a new species and is named Raillietina garrisoni in honor of
the late Dr. P. E. Garrison,
The parasite deserves more than passing notice due to its pos-
sible identity with Davainea madagascariensis (Davaine) of
Garrison, 1911, which was collected at autopsy by Dr. Vernon
L. Andrews from the small intestine of a male adult Filipino
in Manila. According to Joyeux and Baer (1929) Garrison's
material differs in the size of its rostellar hooks and of the cirrus
pouch from the types described under the same name by other
observers, and it is, therefore, likely that it represents another
46,4 Tubangui: Worm Parasites of the Brown Rat 549
species. According to the same authors it is allied to R. cele-
bensis but differs from the latter in the size of its cirrus sac and
in the number of its testes, which characters, it will be recalled,
are the very ones that distinguish R. garrisoni from R. cele-
bensis.
Joyeux and Baer are of the opinion that Garrison's Davainea
madagascariensis and other rare human cestodes are parasites
of wild animals that are accidentally transmitted to man. They
suggest as one way of establishing the identity of these parasites
the systematic collection and determination of the tapeworms
of wild animals that habitually come in close contact with human
beings in countries where such parasites have been recorded.
The survey on which this report is based was, therefore, in line
with the suggestion of the French authors and it is here shown
that there exist important similarities in the morphology of
R. garrisoni and of D. madagascariensis as described by Garri-
son (Table 2). In view of this and in view of the common
occurrence of R. garrisoni in rats, a number of the parasites
of which are transmissible to man, it is quite probable that
Garrisons's tapeworm is identical with this species.
Table 2. — Comparison between Raillietina garrisoni sp. nov. and Davainea
madagascariensis (Davaine) of Garrison, 1911.
D. madagascariensis.
R. garrisoni.
Total length mm_ _
390
Up to 600.
1.60-2.12X1.05-1.40.
0.40-0.80.
0.10-0.15.
90-140.
20-26.
36-50.
0.13-0.18X0.054-0.085.
Normally unilateral; an-
terior.
0.06-0.15 (measured from
mounted specimens) .
1-4; generally 3.
52-80X22-26.
4-6.
Size of terminal gravid segments. _do
Diameter of head do
Diameter of sucker do
Number of rostellar hooks
2.0-2.5X1.0-1.5
0.32-0.40
0. 105-0. 125__
Length of rostellar hooks #,__
Number of testes
23.5-25.2
50
Size of cirrus sac mm__
Position of genital pores
0.12-0. 16X0. 064-0.100.
Normally unilateral; an-
terior.
0.20-0.40
Diameter of uterine egg capsule mm__
Number of eggs per egg capsule
Size of eggs with elongated shell
intact - --fi.~
Length of embryonal hooks #__
1-3 ; generally 2.
50-64X19-23
4-5
Description. — Total length up to 600 millimeters, the maxi-
mum breadth in the region of mature proglottids. Head (fig.
5, 6) subglobular, 0.40 to 0.80 millimeter in diameter; suckers
unarmed, 0.10 to 0.15 millimeter in diameter; rostellum 0.13 to
550 The Philippine Journal of Science 1931
0.18 millimeter in diameter, armed with 90 to 140 hammer-
shaped hooks (fig. 5, a) that are 20 to 26 microns in length and
arranged in two alternating circular rows; rostellum with a
spiny collar, the spines being comma-shaped and averaging about
5 microns long. Neck short, 0.28 to 0.36 millimeter in width.
Segments broader than long except at posterior end where gra-
vid proglottids may be nearly twice as long as wide (fig. 5, d) ;
immature segments 0.08 to 0.17 millimeter long by 0.30 to 0.60
millimeter wide, mature segments 0.43 to 0.65 by 1.40 to 1.65
millimeters, and gravid segments 0.95 to 2.12 by 0.15 to 1.40
millimeters. Genital pores normally unilateral and dextral, sit-
uated near anterior extremity of lateral border of segments.
Main portion of excretory system represented by two pairs
of lateral longitudinal vessels, ventral and dorsal; ventral pair
more lateral in position, larger in diameter and connected in
the posterior part of each segment by transverse canal; dorsal
vessels small and with no transverse canals. Peripheral ner-
vous system represented by a longitudinal nerve on each side,
at middle between ventral excretory vessels and lateral margins
of proglottids. Muscular system feebly developed and arranged
as in other cestodes; consists of minute longitudinal and trans-
verse fibers located immediately beneath cuticle and of longi-
tudinal, transverse and dorsoventral fibers in parenchyma, of
which the longitudinal and dorsoventral ones are most con-
spicuous.
Testes (fig. 5, c) small, roundish, 40 to 50 microns in diame-
ter, confined within parenchyma between excretory vessels, 36
to 50 in number, of which 9 to 15 are on the poral side of the
median line and 26 to 35 aporal. Vas deferens a long, much-
convoluted tube near anterior border of segment, running al-
most transversely from median line to cirrus sac, passing with
corresponding vagina between excretory vessels and ventral to
longitudinal nerve. Cirrus sac distinctly gourd-shaped, 0.13 to
0.18 by 0.054 to 0.085 millimeter in size, extending either trans-
versely or a little obliquely towards cephalic end from genital
pore to longitudinal nerve.
Ovary (fig. 5, c) median, bilobed, each lobe being oval, with
smooth surface and measuring 0.12 to 0.15 by 0.08 to 0.10 milli-
meter. Vagina a narrow canal, posterior to vas deferens and
cirrus pouch, running transversely from median line to genital
pore; before opening into genital pore it is usually slightly
dilated to form a small receptaculum seminis. Near the median
4«,4 Tubangui: Worm Parasites of the Brown Rat 551
0.5 mm
e
Fig. 5. Raillietina garrisoni sp. nov. a, Rostellar hooks; b, scolex ; c. mature segment;
d, gravid segment; e, egg.
line the vagina bends posteriorly and joins the oviduct, forming
a slightly dilated tube, the ootype complex, between ovary and
vitelline gland. Vitelline gland roundish to oval, immediately
posterior to ovary, and measuring 0.09 to 0.13 millimeter across.
552 The Philippine Journal of Science 1931
Uterus at first a simple sac filled with immature ova; in fully
developed gravid segments it breaks down into numerous egg
capsules, each containing 1 to 4, but mostly 3, eggs. Egg cap-
sules (fig. 5, d) 0.06 to 0.15 millimeter in diameter, confined
within excretory vessels, although a few of them may be found
lateral to these canals, and numbering 180 to 200 in anterior
gravid segments and 300 to 400 in elongated posterior gravid
proglottids. Eggs (fig. 5, c) of characteristic shape, the on-
cosphere surrounded by two thin membranes : outer membrane
elongated oval, 52 to 80 by 22 to 26 microns in size; inner mem-
brane usually closely applied around onchosphere, round, 18 to
22 microns in diameter in fresh specimens; between inner and
outer membranes a few connecting strands or fibers are some-
times present ; onchosphere supplied with three pairs of embryo-
nal hooks 4 to 6 microns long.
Specific diagnosis. — Raillietina: Length up to 600 millimeters,
maximum breadth 1.4 millimeters. Head 0.40 to 0.80 millimeter
in diameter; rostellum 0.13 to 0.18 millimeter in diameter, with
90 to 140 hooks 20 to 26 microns long; a spiny collar posterior
to rostellum present, the spines being comma-shaped and about
5 microns long. Suckers unarmed, 0.10 to 0.15 millimeter in
diameter. Genital pores normally unilateral and dextral, near
anterior extremity of lateral border of segments. Cirrus sac
0.13 to 0.18 by 0.054 to 0.085 millimeter in size, extending only
up to nerve. Testes 9 to 15 poral, 26 to 35 aporal, total 36 to
50. Egg capsules 180 to 400, each containing 1 to 4, generally
3, eggs ; found mostly within excretory vessels, but a few lateral
to them.
Location. — Small intestine.
Locality. — Manila, Philippine Islands.
Type specimens. — Philippine Bureau of Science parasitolog-
ical collection, No. 12.
Life history. — Unknown. Probably similar to the mode of
development of most tapeworms, and in particular to other spe-
cies of Raillietina, which utilize as intermediate hosts various
forms of insects.
Prevention. — Due to reasons given above, this tapeworm may
be looked upon with suspicion as one of those parasites of rats
that are transmissible to man. Since the life history has not yet
been worked out, however, no definite prophylactic measures
can be given except to advocate the destruction of rats and mice,
the proper disposal of the stools of infected persons, and the
46,4 Tubangui: Worm Parasites of the Brown Rat 553
practice of all-around cleanliness, by means of which all para-
sitic infestations can be avoided.
References.— 17, 22, 25, 27,
Family HYMENOLEPIDID^E Railliet and Henry, 1909
Subfamily HYMENOLEPIDIN^E Ransom, 1909
Genus HYMENOLEPIS Weinland, 1858
HYMENOLEPIS DIMINUTA (Rudolphi, 1819) Blanchard, 1891. Fig. 6.
Synonyms: T tenia diminuta Rudolphi, 1819; Hymenolepis flavopunc-
tata Weinland, 1858; Tsenia flavomaculata Leuckart, 1863.
This common tapeworm of rats was first reported in man by
Weinland in 1858. Since that time up to 1922, according to
Riley and Shannon (1922), a total of sixty-one cases of human
infestations with this parasite have been recorded from various
parts of the world. To these should be added the one case de-
tected by Schwartz and Tubangui (1922) in a native Filipino,
the twenty Indian cases found by Chandler (1927), and the
single case recently reported by Spindler (1929) from the United
States.
Description. — Strobila composed of 800 to 1,300 proglottids;
length 100 to 600 millimeters, depending upon number of pro-
glottids; maximum width at posterior end in region of gravid
segments, 2.5 to 4.0 millimeters. Head (fig. 6, a) almost glob-
ular, 0.20 to 0.60 millimeter broad ; rostellum rudimentary, pyri-
form, without hooks; suckers globular, near apical portion of
head, 0.08 to 0.16 millimeter in diameter. Neck short. Seg-
ments wider than long; immature segments 0.045 to 0.200 by
0.305 to 0.835 millimeter in size, mature segments 0.238 to 0.380
by 0.084 to 1.670 millimeters, and gravid segments 0.305 to
0.684 by 1.805 to 3.115 millimeters. Posterior border of seg-
ments only slightly wider than anterior borders, for which rea-
son serration of strobila not as marked as in other cestodes.
Genital pores usually unilateral and sinistral, at middle or at
anterior third of lateral margins of proglottids. Main portion
of excretory system consists of two pairs of lateral longitudinal
vessels: a larger ventral pair connected in the posterior part
of each segment by a transverse canal and a smaller dorsal pair
with apparently no cross-connectives ; the terminals of the ven-
tral and dorsal vessels of one side are united in the region of
the head. Muscular system fairly well developed, consisting of
circular and longitudinal subcuticular fibers and another set of
264209 2
554 The Philippine Journal of Science 1931
longitudinal, transverse, and dorsoventral muscle fibers in the
parenchyma.
Normally there are three testes in each mature segment — one
poral and two aporal — arranged, more or less, in a straight
line across segment and separated by ovary (fig. 6, 6). Occa-
sionally this arrangement is reversed; that is, there are two
testes on the poral side of the ovary and one on the aporal side.
Exceptionally, the two aporal testes are placed obliquely or one
behind the other. In some segments, instead of the usual three
testes, there may be only two, or there may be four to six. The
testes are spherical, 0.12 to 0.14 millimeter in diameter. The
vas deferens before entering the cirrus pouch is dilated to form
a prominent seminal vesicle. Cirrus sac 0.17 to 0.30 by 0.02
to 0.04 millimeter in size in mature segment, 0.24 to 0.40 by
0.04 to 0.06 millimeter in gravid segments, extending from
genital pore to or just past excretory vessels; incloses slender,
protrusible cirrus.
Ovary bilobed, 0.35 to 0.40 millimeter across, median, inter-
testicular; surface indented to form small lobules. Vitelline
gland lenticular in shape, immediately postovarial. Shell gland
small, rounded, between ovary and vitelline gland. Recepta-
culum seminis large, prominent, extending transversely from
median line to excretory vessels ; it then becomes narrow in diam-
eter and is continued as the vagina. The latter leads to the
comjmon genital pore, passing ventral and slightly posterior to
the cirrus pouch. Uterus in pregravid segments in the form
of a transversely elongated and apparently solid mass of cells
representing young undeveloped ova; it soon becomes hollowed
out, sending diverticula in all directions, and in the fully devel-
oped state it has the appearance of a sac incompletely divided
by partitions into egg capsules and occupying nearly the entire
space within a gravid segment (fig. 6, c). Mature eggs (fig.
6, d) spherical or slightly oval, the embryo proper or onco-
sphere being surrounded by three membranes, as follows: A
thicker, very faintly radially striated outer membrane, 54 to
86 microns in diameter; a thinner envelope immediately sur-
rounding embryo, oval in shape, 24 by 20 to 40 by 35 microns
in size, often with two polar projections but without filaments
as is the case with the eggs Hymenolepis nana; and an inter-
mediate layer between outer and inner membranes, apparently
composed of albuminous substance and often appearing as two
delicate smooth membranes with intervening space filled by
46,4 Tubangui: Worm Parasites of the Brown Rat 555
Fig. 6. Hymenolepis diminuta. a, Head ; b, mature segment, dorsal view ; c, gravid
ment; d, egg.
granular substance. Embryonal hooks 10 to 16 microns in
length.
Location. — Small intestine.
Life history. — Involves an intermediate host. If ingested by
any of the following insects, the eggs will develop into infectious
larvae known as cysticercoids : Meal moth (Anisopia farinalis),
earwig (Anisolabis annulipes), beetles (Akis spinosa, Scaurus
556 The Philippine Journal of Science 1931
striatus and Tenebrio molitor), cockroaches (Blatta orientalis
and Phyllodromia germanica), and rat fleas (Ceratophyllus fas-
ciatus and Xenopsylla cheopis). The cysticercoids are found
either free in, or encysted in the adipose tissue of, the abdominal
cavities of the above insects. Rats as well as human beings be-
come infected by ingesting these cysticercoids together with any
of the above intermediate hosts.
Prevention. — Consists in the avoidance of rats and mice in
houses, in the destruction of beetles, cockroaches, and other in-
sects that act as intermediate hosts, in the protection of foods
from such insects, and in the proper disposal of the stools of
infected persons.
References.— 8, 14, 25, 27, 33, 35, 40, 48, 49, 51.
HYMENOLEPIS NANA (Siebold, 1852) Blanchard, 1891. Figs. 7 and 8.
Synonyms: Taenia murina Dujardin, 1845; Taenia segyptiaca, Bilharz,
1852; Hymenolepis fratema Stiles, 1906; Hymenolepis longior
Baylis, 1922.
As indicated by its name (nana, or dwarf) one of the dis-
tinguishing characteristics of this cestode is its small size ; hence
it is commonly known as the dwarf tapeworm. It is a common
parasite of rats and mice and of human beings in many parts of
the world, especially in tropical and subtropical countries. Opin-
ion, however, is divided on the identity of the dwarf tapeworm
of rats and mice with the form found in man. Some consider
the two forms as representing one and the same parasite (vide
Woodland, 1924), while others believe that they are distinct
(vide Joyeux, 1925). If the latter opinion should prove to be
true, the rodent parasite would have to be designated as Hyme-
nolepis fratema Stiles, 1906, the older name, Taenia murina
Dujardin, 1845, being preoccupied and, therefore, not available.
The designation nana would then apply only to the human form.
The present writer believes with Stiles (1906) that " from a
standpoint of prevention they should at present be considered
as identical," which opinion has been justified by the successful
cross-infection experiments of Saeki (1920) and Woodland
(1924) as well as by the recent epidemiological observations of
Chandler (1927). The latter investigator concluded from his
observations that rats are an important epidemiological factor
in the dissemination of H. nana, for he found the distribution of
the parasite in human beings in India to correspond very closely
with that of another rat-borne disease ; namely, bubonic plague.
The occurrence of this parasite in human beings in the Phil-
ippines has been recorded by Riley (1919), who found it in a
46,4
Tubangui: Worm Parasites of the Brown Rat 557
faecal sample obtained from an American Bohemian child resid-
ing in Zamboanga, Mindanao, and forwarded to him by Dr. A.
F. Coutant. The child was one of a family of five and it appears
from the data furnished by the sender that the other members
of the family were similarly infested with the worm in question.
Among the files of the Bureau of Science for 1928 on the results
of the routine examination of faecal specimens submitted by the
Philippine Health Service for evidences of intestinal parasitism,
there is also an unpublished record of its presence in a young
Chinese boy living in Manila. In Philippine rats, on the other
hand, this is the first report of its occurrence, and it seems that
it is rare in these animals, for it was found in only 1.7 per cent
of the total number of rats examined.
Description. — Strobila composed of
96 to 840 proglottids; length 5 to 90
millimeters, depending upon number
of segments ; maximum width 0.20 to
0.90 millimeter, near posterior end.
Head (fig. 8, a) subglobular, 0.13 to
0.48 millimeter in diameter; suckers
globular, 0.07 to 0.15 millimeter in
diameter; rostellum well developed,
freely movable, armed near its anter-
ior end with 20 to 30 characteristic
hooks (fig. 7, b) ; latter 14 to 18 mi-
crons in length, with curved dorsal
root directed anteriorly on rostellum
and, directed posteriorly, a thick ven-
tral root about equal in length to a
sharp pointed prong with which it
forms a sort of fork. Neck slender,
0.08 to 0.10 millimeter in length by
0.08 to 0.30 millimeter in width. An-
terior segments very short; follow-
ing segments increase in length and
breadth but remain broader than
long; most posterior segments, how-
ever, may be occasionally stretched
and be as long as wide or even longer
than wide. Measurements on Philippine material as follows:
Immature segments 0.02 to 0.03 millimeter long by 0.14 to 0.17
millimeter wide, mature segments 0.04 to 0.08 by 0.17 to 0.32
millimeter, gravid segments 0.08 to 0.12 by 0.30 to 0.37 millime-
Fig. 7. Hymenolepis nana, a, En-
tire worm (from Ransom, 1904) ; bt
rostellar hooks.
558 The Philippine Journal of Science 1931
ter. Genital pores generally all on left side, near anterior bor-
der of segments.
Main portion of excretory system consists of two pairs of
lateral longitudinal excretory vessels : a small dorsal pair and a
larger ventral pair of vessels, the latter united in the posterior
portion of each segment by a transverse canal; ventral and
dorsal vessels of one side united in the region of the scolex
and form an anastomosis at the base of the rostellum. Peri-
pheral nervous system represented by a pair of longitudinal
nerves, one on each side of strobila, lateral to excretory vessels.
Muscular system weakly developed, consisting of outer circular
and inner longitudinal subcuticular fibers and of longitudinal
fibers in parenchyma ; transverse and dorsoventral parenchymal
fibers may also be present, but very few and weakly developed.
Three testes in each mature segment (fig. 8, 6), normally one
on left and two on right side of median line and usually ar-
ranged in more or less straight transverse line at posterior por-
tion of proglottids; the arrangement, position, and number of
these organs, however, are liable to variation as in Hymeno-
lepis diminuta; they are globular, 28 to 34 microns in diameter.
Vas deferens a slender canal for the most part; before enter-
ing cirrus pouch it may be dilated to form a small seminal
reservoir; within cirrus pouch it may also be enlarged to form
a seminal vesicle. Cirrus pouch club-shaped, 0.065 to 0.072 by
0.018 to 0.021 millimeter in size, its long axis directed trans-
versely or sometimes obliquely forwards from genital pore to
excretory vessels, passing dorsal to longitudinal nerve.
Ovary transversely elongated, bilobed, 0.10 to 0.12 millimeter
across, lying ventral to testes. Vitelline gland rounded to oval
in shape, immediately postovarial. Shell gland very small, be-
tween ovary and vitelline gland. Receptaculum seminis large,
prominent, extending transversely from median line to excre-
tory vessels ; it then becomes narrow in diameter and is contin-
ued as the vagina. Latter leads to common genital pore, pas-
sing between cirrus pouch and excretory vessels and nerve.
Uterus at first a transversely elongated cellular mass in front of
ovary; it soon hollows out and assumes in the oldest segments
the form of a sac containing many inf oldings or incomplete par-
titions (fig. 8, c) ; it is more or less completely filled with eggs
numbering 80 to 180 in each gravid segment. Eggs (fig. 8, d)
oval or globular, with two distinct membranes separated by an
intervening space containing a finely granular transparent sub-
46,4 Tubangui: Worm Parasites of the Brown Rat 559
Fig. 8.
Hymenolepw nana, a, Head ; b, mature segment, ventral view ; c, gravid segment ;
d, egg.
stance; outer egg membrane, according to various authors, 30
to 60 microns in diameter; inner membrane 16 to 34 microns in
diameter, usually with more or less conspicuous mammillate pro-
jection at each pole and filamentous appendages; embryonal
hooks 10 to 14 microns long. (Measurements of eggs of Phil-
ippine material as follow: Outer membrane 45 to 60 by 34 to
51 microns, inner membrane 30 to 34 by 23.5 to 27.2 microns.)
Location. — Small intestine.
Life history. — This parasite is unique among the other ces-
todes in that it has a one-host life-cycle; that is, it is capable
of completing its development from egg to adult in a single in-
dividual host. This peculiar life history was first demonstrated
by Grassi (1887) and has subsequently been confirmed by the
more recent studies of Joyeux (1920), Woodland (1924), and
others.
The mature eggs (onchospheres) are discharged with the
faeces of an infested animal. If swallowed by a proper host
560 The Philippine Journal of Science 1931
(for example, a rat), the inclosed embryos become free in the
intestinal tract and develop into cysticercoids within the intes-
tinal villi. The cysticercoids then reenter the alimentary canal
where they grow into adult tapeworms.
Prevention. — Avoid rats and mice in houses; keep foods out
of the reach of rats and mice, especially foods that are eaten
raw, or after cooking are kept for some time before being
eaten ; avoid introducing into the mouth dirty and unnecessary
objects that are apt to be contaminated with the eggs of the
parasite. Infested persons should observe strict personal clean-
liness, especially after defecation and their stools should be prop-
erly disposed of.
References.— 8, 14, 18, 23, 24, 25, 27, 33, 36, 48, 49, 51, 58,
59.
Phylum NEMATHELMINTHES Vogt (quoted by
Carus, 1863)
Class NEMATODA Rudolphi, emend. Diesing, 1861
Order EUNEMATODA Ward, 1916
Superfamily RHABDIASOIDEA Railliet, 1916
Family RHABDIASIDJE Railliet, 1915
Genus STRONGYLOIDES Grassi, 1879
STRONGYLOIDES RATTI Sandground, 1925, fig. 9.
Synonym: Strongyloses papillosus (Wedl, 1856) Hall, 1916.
This minute worm was found in scrapings from the mucous
membrane of the small intestine of 74 per cent of the rats ex-
amined. In a large number of the cases it was associated with
Nippostrongylus muris. As indicated by Sandground (1925),
it may be distinguished from S. papillosus (Wedl, 1856) of
sheep, goats, and rabbits, with which it has been confused, by
its smaller size, the finer striations of its cuticula, and the course
of its ovaries.
Description. — Parasitic generation, represented by females,
2.20 to 2.75 millimeters long by 30 to 35 microns thick. Body
filiform, attenuated anteriorly; posterior end behind anus sud-
denly tapers into a short pointed tail. Cuticle finely striate.
Mouth surrounded by three minute papillae; leads directly to
cesophagus. (Esophagus 0.70 to 0.78 millimeter long, gradually
increasing in diameter posteriorly. Excretory pore 0.10 to 0.12
46,4 Tubangui: Worm Parasites of the Brown Rat 561
OV'
oe
M
S
CM
O
€t
V
OV
Fig. 9. Strongyloides ratti, entire worm, a, Anus ; oe, oesophagus ; ov, ovary ; v, vulva.
millimeter from anterior end. Nerve ring immediately in front
of excretory pore. Anus 42 to 45 microns from posterior end.
Vulva with prominent lips, 1.70 to 1.82 millimeters from ante-
rior end. Ovaries directly recurrent, their bends being close
to oesophageal and anal ends of digestive tract; each is continued
as oviduct, then as uterus, so that uteri are divergent. Eggs
few in number (maximum 10 or 11 in both uteri), 51 to 56
by 27 to 29 microns in size (according to Sandground, 47 to 52
by 28 to 31 microns) ; they contain larvae at deposition.
Location. — Small intestine.
Life history. — As shown by Sandground (1926), the life his-
tory is very similar to that of Strongyloides stercoralis of man.
The eggs hatch while still in the small intestine of the host
562 The Philippine Journal of Science 1931
and the liberated, actively motile, rhabditiform embryos are
passed with the feces. These may either develop immediate-
ly into filariform larvae that are capable of infesting new hosts
or become mature free-living males and females that copulate
and produce eggs, from which free-living rhabditiform larvae
are hatched. The latter are then transformed into infective fila-
riform larvae. Infestation is usually through the skin, the fila-
riform larvae being capable of boring through the integument
of the host.
References.— -7, 19, 38, 63.
Superfamily TRICHUROIDEA Railliet, 1916
Family TRICHOSOMOIDIDJE Yorke and Maplestone, 1926
Subfamily TRICHOSOMOIDIN/E Hall, 1916
Genus TRICHOSOMOIDES Railliet, 1895
TRICHOSOMOIDES CRASSICAUDA (Bellingham, 1840) Railliet, 1895. Fi*. 10.
Synonyms: Trichosoma crassicauda Bellingham, 1840; Triehosoma
muris decumani Rayer, 1843.
Description. — Marked sexual dimorphism : male much smaller
than, and usually parasitic in vagina or uterus of, female (fig.
10, a). Anus posteroterminal in both sexes.
Male 1.60 to 5.20 millimeters in length by 19 to 40 microns in
maximum width, according to various authors. Body thread-
like, not distinctly divided into slender anterior and enlarged
posterior portions. Cuticle very finely striated transversely.
Anterior end (fig. 10, b) with terminal stylet and prepucelike
cuticular sheath, according to Thomas (1924). (Esophagus 0.70
to 1.28 millimeters long or about one-half to one-third of total
body length. Testis single, tubular, originating from anterior
region of body and extending to near posterior end, where it
is transformed into a small seminal vesicle. Spicule, bursa, or
copulatory organs of any sort absent.
Female 10.5 to 14.6 millimeters in length by 0.175 to 0.200
millimeter in maximum thickness near posterior end. Body cov-
ered with transverse cuticular ridges except at extreme anterior
end ; it is divided into a slender anterior portion, corresponding
to length of oesophagus, and into a thicker posterior portion oc-
cupied by intestine and reproductive organs. Head rounded, 20
46,4 Tubangui: Worm Parasites of the Brown Rat 563
Fig. 10. Trichosomoides crassicauda. a, Mature female with male in uterus (after Hall,
1916), m, male worm; v, vulva; b, anterior end of mature male (after Thomas, 1924) ;
c, egg.
to 22 microns in diameter. Mouth simple, minute. (Esopha-
gus a capillary tube, 1.50 to 1.90 millimeters long, which is equal
to between one-sixth and one-eighth of total body length; most
of the anterior portion of oesophagus apparently free of surround-
ing cells, the rest passing through a chain of large oesophageal
cells. Vulva ventral, immediately behind oesophageal termina-
tion. Vagina long, thin-walled, directed posteriorly, distinguised
from uterus by presence of dark brown eggs. Uterus reaches to
near posterior end of body. Eggs (fig. 10, c) generally oval
but may be subspherical or cylindrical, thickshelled, plugged at
both poles, embryonated at deposition, 61.2 to 72.0 by 25.0 to
564 The Philippine Journal of Science 1931
56.0 microns in size. They are colorless in the uterus, but in the
vagina they become dark brown.
Location. — Urinary bladder ; also renal pelvis and ureters.
Life history. — Simple and direct. If ingested by the proper
host, the eggs, which contain well-developed embryos when ovi-
posited and which are passed out with the urine of an infected
animal, hatch in the stomach of the host. The newly-hatched
larvae measure 264 to 390 microns in length by 10 to 16 microns
in maximum width and are provided with a terminal stylet and
a prepucelike fold at the anterior end. After boring out through
the wall of the digestive tract these larvae enter the blood stream
and are carried to the heart by way of the portal system. Ac-
cording to Yokogawa (1921), they have to pass through the
lungs before they can establish themselves in their normal hab-
itat. The experiments of Thomas (1924), however, do not in-
dicate that passage through the lungs is absolutely essential in
the development of this parasite in the same sense that Ascaris
larvae, for example, must go through these organs before they
can become adults. According to Thomas, the larvae of Tricho-
somoides crassicavda are dispersed by the circulatory system
to different parts of the body, but only those that become lodged
in the urinary tract are able to complete their development.
The adult state is reached in three to six weeks after the in-
gestion of the eggs. Copulation takes place at any point in
the urinary tract. The male enters the vagina of the female
and may either remain there permanently or wander out again.
References.— 19, 52, 61, 63.
Family TRICHURIDJE Railliet, 1915
Subfamily CAPILLARIIN^E Railliet, 1915
Genus HEPATICOLA Hall, 1916
HEPATICOLA HEPATICA (Bancroft, 1893) Hall, 1916. Fig. 11.
This appears to be one of the commonest parasites of the
brown rat in the Philippines, about 90 per cent of the rat
livers examined showing the presence of irregular white or
yellowish spots that mark the presence of the worm's eggs.
It is at the same time one of those helminths that are able to
establish themselves in a variety of hosts other than rats and
mice. It has been reported from the European hare (Lepus
europus), the rabbit, and the prairie dog (Cynomys ludovicia-
nus) . The guinea pig, dog, and monkey are also susceptible to
46,4 Tubangui: Worm Parasites of the Brown Rat 565
it. In man the first and, up to the present time, the only re-
port of its occurrence is that by Dive and Lafrenais (1924),
who recovered the parasite from a
British soldier who lived for three
years in India. At autopsy the
subject presented a liver abscess,
in the proximity of which were
masses of the parasite's eggs; the
worms themselves were found in
the periphery of the abscess.
Description. — Body capillary di-
vided into anterior oesophageal and
posterior portions. Cuticle del-
icately striate, apparently without
bacillary band. Mouth simple.
Worms, both male and female, 40
to 50 millimeters long.
Male 28 microns thick at poster-
ior end ; anterior and posterior por-
tions of body about equal in length.
Spicule absent, but represented by
membranous sheath prolonged
from posterior extremity.
Female 100 to 120 microns thick
at middle of body and 65 microns
at tail. Anterior portion of body
about half as long as posterior
portion. Vulva (fig. 11, a) prom-
inent, 6 to 7 millimeters from an-
terior end, opening at level of
posterior oesophageal region. Tail
very short, blunt and conical. Ovi-
parous; eggs lemon-shaped, double
walled, 54 to 58 by 32 to 34 mi-
crons in size, plugged at each pole
(fig. 11, &) ; outer eggshell striate,
inner shell homogeneous.
Location. — Liver.
Life history. — The life history is simple and direct; that is,
it does not involve any intermediate host. The eggs, as en-
countered in the liver of recently dead rats, are nonsegmented
or in the very early stage of segmentation, and are not infectious.
Fig. 11. Hepaticola hepatica. a, An-
terior end of mature female (after
Nishigori, from Yorke and Maple-
stone, 1926); oe, oesophagus; v,
vulva ; b, egg.
566 The Philippine Journal of Science 1931
Their development to the infectious embryonated stage is quite
slow, taking about five to six months according to Bancroft and
to Railliet, and only twenty-three days according to Momma
(1930). If mature eggs are ingested by a proper host, hatch-
ing takes place in the small intestine and the newly liberated
larvae, after penetrating through the intestinal wall, reach the
liver by way of the circulatory system, according to Fulleborn
(1924). According to Nishigori (1925) and Asada (1925),
they pass through the intestinal wall into the abdominal cavity,
from which they make their way into the liver through the
surface of this organ. In any case, the larvae, after reaching
the liver, stay there to complete their development. Larvae may
sometimes be carried to the lungs and other organs, in which
case they do not become mature and sooner or later die.
In connection with this mode of development it is not yet
clear how the eggs are discharged from the body and how
they are transmitted from one host to another. According to
Railliet (1892) and others who state that they have seen the
eggs in the faeces, it is presumed that they escape through the
intestinal tract from the liver through the bile duct. On the
other hand, according to Bancroft (1893), Weidman (1925),
and others who have failed to detect their presence in the fae-
ces of infested animals, the belief is that the transmission of
the eggs probably depends upon the cannibalistic habit of the
host animals. This, however, could hardly be considered in
the case of the human infestation recorded above. Even with
rats direct infestation through cannibalism is possible only if
a rat will devour another rat (infested) that has been dead for
several months and in which the eggs have had time to develop
into the infective stage. Indirectly, however, cannibalism may
play a distinct role in the spread and propagation of the pa-
rasite if it could be shown that the immature eggs, as found
in the liver, will still continue their development after passing
through the digestive tract of a rat. In this connection the
recent observations of Momma (1930) and Shorb (1931) are
interesting. These authors cultured eggs derived from the fae-
ces of flies and cats that had been fed on infested rat livers and
found that they developed normally to the infective stage. It
may, therefore, be deduced that the eggs of Hepaticola hepa-
tica are disseminated through the natural decomposition and
disintegration of the dead bodies of infested animals and through
the capture and ingestion of infested rats and mice by their own
46, 4 Tubangui: Worm Parasites of the Brown Rat 567
kind, or by cats, and other rat-preying animals. According to
Momma, flies may play a role in the dispersal of the ova since
they are often seen in large numbers around the decomposing
bodies of dead rats.
Prevention. — Avoid rats and mice in houses ; the dead bodies
of these animals should not be allowed to decompose in the
open, but should be buried deeply in the ground or burned;
protect foods from rodents and from flies.
References.— 2, 3, 12, 14, 15, 19, 28, 29, 32, 45, 51, 57, 63.
Superfamily STRONGYLOIDEA Weinland, 1858; Hall, 1916
Family TRICHOSTRONGYLID^ Leiper, 1912
Subfamily HELIGMOSOMIN-^E Travassos, 1914
Genus NIPPOSTRONGYLUS Lane, 1923
NIPPOSTRONGYLUS MURIS (Yokogawa, 1920) Lane, 1923. Fig. 12.
Synonym: Heligmosomum muris Yokogawa, 1920.
Description. — Body small, filiform, coiled, blood red in color
when fresh. Cervical ate absent, but cuticle inflated in head re-
gion (fig. 12, a) ; length of cuticular expansion 0.058 to 0.063
millimeter. Cuticle with ten longitudinal ridges originating be-
hind inflated area; transverse striation of cuticle evident on
these ridges. Mouth simple, leading into small buccal cavity.
(Esophagus 0.30 to 0.40 millimeter long. Nerve ring 0.20 to
0.23 millimeter from anterior end. Excretory pore a short
distance in front of nerve ring. Cervical papillae lacking.
Male 3.2 to 3.5 millimeters long by 0.08 millimeter in max-
imum thickness at middle of body. Bursa well developed, with
conspicuous asymmetrical lateral lobes and rays and small dor-
sal lobe (fig. 12, b). Right lobe larger, at least longer, than
left lobe, its supporting rays differing from those of opposite
side: ventroventral ray small, slender, widely separated from
lateroventral which is also thin but longer; externolateral and
mediolateral thick and close together except at their tips; pos-
terolateral small and delicate; externodorsal on both sides thin
and slender, arising at slightly higher level from common trunk
with dorsal ray. In the left lateral bursal lobe, the ventro-
ventral, lateroventral, externolateral and mediolateral rays are
almost similar in form, being long and thin; posterolateral
thicker and curved dorsally, ending in a conical tip. Dorsal
ray bifurcate at its tip, each limb ending in two or three di-
gitations. Spicules yellowish in color, equal, filiform, 0.44 to
568
The Philippine Journal of Science
1931
a b
Fig. 12. Nippostrongylus muris. a, Anterior end, lateral view; b, bursa, dorsal view.
0.50 millimeter in length by 6 to 7 microns in maximum width
at their proximal ends, with sickle-shaped extremities that are
usually united together. Gubernaculum colorless, 44 to 46 mi-
crons long.
Female 4.0 to 4.6 millimeters in length by 0.135 millimeter
in maximum thickness at middle of body. Posterior end be-
hind vulva reduced abruptly in diameter ending in a short,
curved, conical tail; in contracted specimens this region of the
body may appear swollen and bell-shaped due to the invagina-
tion of the cuticle which carries with it the anus and the vulva.
Anus about 32 microns from tip of tail. Vulva in front of
anus, about 80 microns from tip of tail ; vagina muscular, sep-
arated from uterus by ovejector; uterus short, modified an-
teriorly into receptaculum seminis; ovary long, with short an-
terior loop. Eggs few in number, thin shelled, segmented at
deposition, 58 to 60 by 30 to 32 microns in size.
Location. — Small intestine.
Life history. — The life history of this nematode has been
worked out by Yokogawa (1922) . When passed out in the faeces
of the host, the eggs are in various stages of segmentation. Un-
der favorable conditions their development is continued outside
and hatching takes place after about twenty to twenty-four hours.
46,4 Tubangui: Worm Parasites of the Brown Rat 569
The newly hatched larvae attain the infective stage after about
five days. The infection of new hosts is accomplished, as in
the case of hookworms, by the larvae entering the body either
through the skin or by way of the mouth, the former method
having been shown to be more effective. After passing through
the lungs the larvae settle down in the intestine where they reach
sexual maturity in seven to ten days after infestation.
References.— 26, 60, 62, 63.
Superfamily OXYUROIDEA Railliet, 1916
Family OXYURID^E Cobbold, 1864
Subfamily SYPHACIIN^ Railliet, 1916
Genus SYPHACIA Seurat, 1916
SYPHACIA OBVELATA (Rudolphi, 1802) Seurat, 1916. Fifif. 13.
This parasite is listed by Shipley (1908) and by Stiles and
Hassall (1910) among the nematodes reported from Mus norve-
gicus. As already stated, it was not encountered in the pre-
sent survey, but attention is called to it in view of its recorded
occurrence in the Philippine Islands by Riley (1919) who iden-
tified it from specimens found in a sample of human stools ob-
tained from an American Bohemian child residing in Zamboan-
ga, Mindanao, and forwarded to him by Dr. Albert F. Coutant.
The following description is mostly adopted from Hall (1916).
Description. — Body elongate, fusiform. Cuticle transversely
striate, not dilated in head region. Two small cervical ate pre-
sent (fig. 13, a). Mouth bounded by three lips, each bearing a
median papilla on its outer face; mouth cavity simple. (Eso-
phagus club-shaped with a posterior bulb containing a valvular
apparatus and separated from the rest by a constriction. Ex-
cretory pore a little posterior of level of oesophageal bulb.
Male (fig. 13, b) 1.3 millimeters long by 115 microns thick,
with two or three cuticular "mamelons" on ventral surface.
Posterior extremity coiled in a spiral and ending in a long
pointed tail. Narrow caudal alse present, limited to first part of
tail, supported by two pairs of preanal and one pair of postanal
pedunculated papillae (fig. 13, c). Spicule simple, slightly
curved, 85 microns long by 7 microns thick at base; guberna-
culum shaped like a ploughshare, 37 microns long, directed trans-
versely posterior of spicule. Cloacal aperture 210 microns from
tip of tail ; posterior lip of aperture with a small chitinous hook
that may be of use in copulation.
2642C9-- — 3
570
The Philippine Journal of Science
1931
Fig. 13. Syphacia obvelata. a, Anterior end of female, lateral view ; b, male, lateral view ;
c, posterior end of male, ventral view ; d, posterior end of female, lateral view ; e\ ege.
(All from Yorke and Maplestone, 1926.)
Female 3.5 to 5.7 millimeters long by 115 to 215 microns
thick. Body terminates in a long, narrow tip posteriorly (fig.
13, d.) (Esophagus, exclusive of bulb, 255 to 330 microns long
by 50 to 70 microns thick ; oesophageal bulb 85 to 100 by 75 to
110 microns in size. Nerve ring 100 to 130 microns from an-
terior end. Anus 515 to 705 microns from tip of tail. Vulva
prominent, behind excretory pore, situated on conical cuticular
prominence 540 to 740 microns posterior of head. Vagina ex-
tends posteriorly from vulva, elongate, about 170 microns long.
Uterine branches do not extend posterior of anus. Eggs 110
to 142 by 30 to 40 microns in size, nonembryonated at time of
oviposition (fig. 13, e).
Location. — Caecum and large intestine.
Life history. — Unknown. Probably similar to that of closely
related nematodes, such as, Enterobius vermicularis, the human
pin worm, the life history of which is simple and direct.
Prevention. — Taking for granted that the life history of this
parasite is simple and direct, the preventive measures that sug-
gest themselves are the observance of personal cleanliness, es-
46,4 Tubangui: Worm Parasites of the Brown Rat 571
pecially after defecation, the proper disposal of the stools of in-
fected individuals, the destruction of rats and mice, and the pro-
tection of foods from the droppings of these animals.
References.— U, 19, 34, 42, 44, 50, 51, 63.
Family HETERAKIDJE Railliet and Henry, 1914
Subfamily HETERAKIN^E Railliet and Henry, 1912
Genus HETERAKIS Dujardin, 1845
HETERAKIS SPUMOSA Schneider, 1866. Fig. 14.
Synonym: Ganguleterakis gangula Lane, 1914.
Descriptidn. — Body small, tapering slightly towards the an-
terior end. Cuticle with fine longitudinal and transverse stria-
tions and with lateral flanges in oesophageal region (fig. 14, a) .
Head 70 to 75 microns in diameter. Mouth with three subequal
lips, each lip carrying two lateral papillae. (Esophagus 0.75 to
0.83 millimeter long, subcylindrical, terminating in a well-de-
veloped bulb; latter 0.15 to 0.17 millimeter in diameter, pro-
vided with a valvular apparatus. Distance from anterior end to
nerve ring 0.22 to 0.24 millimeter; to excretory pore 0.29 to
0.31 millimeter; to cervical papillae 0.30 to 0.34 millimeter.
Male 6.0 to 7.4 millimeters in length by 0.25 millimeter in
maximum thickness. Tail short and sharply pointed. Caudal
alae well developed, provided with ten pairs of papillae grouped
as follows (fig. 14, c) : an anterior group of two pairs of ven-
tral papillae lateral to genital sucker, a middle group of two
pairs of ventral and three pairs of lateral papillae in the cloacal
region, and a posterior group of three pairs of lateral papillae
near tip of tail. The anterior group of papillae are all slender ;
in the middle group the ventral pairs are short and knobby,
while the lateral pairs vary in size and appearance, the first pair
being the largest, the second pair thick but short, and the last
pair longer but slender; the posterior group of papillae are rel-
atively small, the middle pair being the largest among them.
Genital sucker slightly oval transversely, pedunculate, with a
strong chitinous rim interrupted posteriorly by a papilliform
projection; average size 0.076 by 0.087 millimeter and about
0.15 millimeter from cloacal opening. Spicules subequal, taper-
ing distally, 0.280 to 0.315 millimeter in length. Gubernaculum
absent. Cloacal opening 0.29 to 0.32 millimeter from tip of tail.
Female 7.8 to 9.5 millimeters in length by 0.30 to 0.32 milli-
meter in maximum thickness. Tail long and acutely pointed
(fig. 14, &). Anus 0.58 to 0.64 millimeter from tip of tail.
572
The Philippine Journal of Science
1931
Fig. 14.
Heterakis spumosa. a, Anterior end of female, ventral view ; b, posterior end of
female, lateral view ; c, posterior end of male, ventral view.
Vulva slightly posterior of middle of body length. Vagina mus-
cular, is at first directed anteriorly, then bends posteriorly and
divides into anterior and posterior uterine branches. Ovaries in
numerous transverse coils in anterior and posterior end of body.
Eggs oval, thick shelled, in the early stage of segmentation at
deposition, 56 to 65 by 38 to 40 microns in size; shell about 4
microns in thickness.
Location. — Large intestine (caecum).
Life history. — Not worked out, but possibly similar to that of
Heterakis gattinse of poultry, in which case it is simple and di-
rect. Briefly the life history of H. gallinse is as follows: The
eggs are passed outside with the faeces of the host. Under
46, 4 Tubangui: Worm Parasites of the Brown Rat 573
favorable conditions of temperature and moisture, the egg be-
comes embryonated; that is, a larva is developed inside each
egg, and is then infective. If the egg is swallowed by a proper
host, hatching takes place in the intestine and the liberated
larva soon settles down in the caecum to grow into an adult.
The larvae do not wander into the lungs as is the case with the
larvae of Ascaris.
References. — 19, 63.
Superfamily SPIRUROIDEA Railliet and Henry, 1915
Family SPIRURIDiE Oerley, 1885
Subfamily SPIROXYIN^ Baylis and Lane, 1920
Genus PROTOSPIRURA Seurat, 1914
PROTOSPIRURA MURICOLA Gedoelst, 1916. Fig. 15.
This is possibly the small Ascaris which Schobl (1913) has
observed as being not uncommon in the intestine of Philippine
rats. Its normal habitat is the stomach, but after the death of
the host it often migrates into the small intestine.
The specimens at hand differ greatly among themselves in
size, some females in particular being almost twice as large as
other females. In the beginning it was thought that the collec-
tion represented two species, but it was later revealed that out-
side of size there were no other morphological differences.
Description. — Body relatively large, regularly attenuated an-
teriorly. Cuticle transversely striated. Mouth (fig. 15, 6)
with two large lateral lips, each divided into three lobes, of
which the middle is larger; each lobe bears two cuticular pro-
jections, but no teeth. There are five pairs of head papillae;
namely, one large pair of subventral, a smaller pair of subme-
dian, the dorsal homologues of these, and a minute pair of lateral
papillae. Pharynx (fig. 15, a) prominent, laterally compressed,
with thick chitinous wall. GEsophagus very elongate, subcylin-
drical, slightly constricted in region of nerve ring, separated
from intestine by valvular apparatus. Cervical papillae not
prominent, in front of nerve ring. Excretory pore ventral,
behind nerve ring.
Male 25 to 30 millimeters in length by 0.80 millimeter in
maximum thickness at middle of body. Average length of
pharynx 0.09 millimeter; of oesophagus 6.20 millimeters. Dis-
tance from anterior end to cervical papillae 0.32 to 0.35 milli-
meter; to nerve ring 0.38 to 0.41 millimeter; to excretory pore
574
The Philippine Journal of Science
1931
Fig. 15. Protospirura muricola. a, Anterior end of female, ventral view; b, mouth, an-
terior view ; c, posterior end of male, lateral view ; d, posterior end of female, lateral
view.
0.43 to 0.45 millimeter. Caudal end of body conical, spiral,
carrying moderately developed symmetrical bursa (fig. 15, c).
Latter with cuticular oblong markings and usually supported by
nine pairs of pedunculated papillae, of which four pairs are
larger and preanal in position and five pairs smaller and post-
anal. Sometimes an extra pair of minute papillse is present near
46, 4 Tubangui: Worm Parasites of the Brown Rat 575
tip of tail. Both spicules bent, with enlarged proximal extrem-
ities and pointed distal ends, but unequal in size and structure;
left spicule spongy, larger, 0.40 to 0.43 millimeter in length by
0.058 to 0.060 millimeter in maximum thickness at proximal
end; right spicule hollow, 0.36 to 0.39 by 0.041 to 0.042 milli-
meter in size. Gubernaculum small, slender, 0.10 millimeter
long. Average distance from tip of tail to cloacal opening 0.42
millimeter.
Female 35 to 52 millimeters in length by 1.20 millimeters in
maximum thickness at middle of body. Pharynx 0.10 to 0.13
millimeter, oesophagus 6.40 to 7.90 millimeters long. Distance
from anterior end to cervical papillae 0.33 to 0.40 millimeter;
to nerve ring 0.38 to 0.47 millimeter; to excretory pore 0.48 to
0.70 millimeter. Caudal end of body bluntly conical (fig. 15,
d). Anus 0.40 to 0.42 millimeter from posterior end. Vulva
a short distance in front of middle of body length. Uteri di-
vergent, anterior uterus reaching anteriorly to almost as far as
oesophago-intestinal junction and the posterior uterus extending
to a short distance in front of anus. Eggs oval, embryonated
at deposition, thick shelled, 50 to 57 by 38 to 44 microns in
size.
Location. — Stomach.
Life history. — Probably similar to that of Protospirura muris
(Gmelin, 1790) and of P. Columbiana Cram, 1926, in which in-
termediate hosts are involved. Cram gives the life history of
P. columbiana as follows : If the embryonated eggs of the pa-
rasite are fed to cockroaches (Phyllodromia germanica), the
liberated larvae find their way to the body cavity, where they
begin to encyst in about a month after feeding. The cysts, how-
ever, are not infective at this time. After forty-one days they
appear to have reached that stage and if fed to rats the encysted
larvae are capable of pursuing further development in the
stomach of the latter. They become fully grown and mature
one hundred fifteen days after the feeding of the final host.
References.— 6, 9, 19, 40, 63.
Subfamily GONGYLONEMINi€ Hall, 1916
Genus GONGYLONEMA Moliit, 1857
GONGYLONEMA NEOPLASTICUM (Fibber and Ditlevsen, 1914) Ransom and Hall, 1916
Fiirs. 16 and 17.
Synonym: Spiroptera neoplastica Fibiger and Ditlevsen, 1914.
Two species of the genus Gongylonema have been reported
from rats ; namely, G. neoplasticum and G. orientale Yokogawa,
576 The Philippine Journal of Science 1931
1925. They are said to differ from each other in the following
respects: Morphologically, in total size, length of oesophagus,
spicules and vas deferens, structure of the spermatozoa, size
of the eggs, etc ; biologically, in the time necessary for the sexes
to reach maturity in experimental infestations. In view, how-
ever, of the observations of Seurat (1916) and Baylis (1925)
on the degree of morphological variations exhibited by members
of the genus, it is not unlikely that the two rodent parasites
are identical. Baylis even goes further in suspecting that G.
neoplasticum is similar to G. pulchrum of the pig, between which
the differences are much greater and, therefore, more apparent.
This, however, could hardly be the case, for, if it were so, it
would be difficult to explain why a parasite that is so common
in rats has not yet been reported in Philippine domesticated
animals. The writer has looked for G. ptdchrum with uni-
formly negative results in swine, sheep, goats, and cattle.
This parasite has received considerable attention due to the
report of Fibiger and Ditlevsen (1914) that it is instrumental
in the production of carcinomatous growths in rats. In the
present survey this possible role of the parasite was kept con-
stantly in mind, but of the rats found harboring it not one pre-
sented a gastric tumor. The condition must be rare in Philip-
pine rats, for Schobl (1913), who examined tens of thousands
of these animals in connection with plague, records only one
case of tumor located on the large curvature of the stomach.
No determination was made as to the possible origin of the new
growth.
Description. — Body long, slender, threadlike, terminating in
a blunt cone anteriorly. Cuticle transversely striate; bears in
cephalic and oesophageal regions more or less globular, egg-
shaped or sausage-shaped cuticular plaques or bosses, of variable
size and arranged irregularly in longitudinal rows on body sur-
face (fig. 16). Lateral bands present, extending on both sides
throughout body length except at most anterior and most pos-
terior regions. Cervical papillae inconspicuous, in front of nerve
ring. Excretory pore ventral, behind nerve ring. Mouth small,
surrounded by four very inconspicuous lips ; buccal rim 0.02 to
0.03 millimeter in diameter. (Esophagus very long, in two
parts — anterior muscular and posterior glandular — separated
from intestine by constriction and intestinal valves.
Male 11.0 to 12.0 millimeters in length by 0.20 millimeter in
maximum thickness at middle of body. Pharynx 0.05 millime-
46,4 Tubangui: Worm Parasites of the Brown Rat 577
ter long. Anterior portion of oesophagus 0.4 millimeter long,
posterior portion 2.4 millimeters; total length of oesophagus,
therefore, about one-fourth of total body
length. Distance from anterior end of
worm; to cervical papillae 0.13 millimeter,
to nerve ring 0.22 millimeter, to excretory-
pore 0.34 millimeter. Tail (fig. 17, a)
slightly twisted on its long axis, provided
with asymmetrical alse, the left wing being
usually longer than the right. Eight pairs
of pedunculated caudal papillae present,
of which four pairs are slightly larger
and preanal and four pairs postanal; last
postanal pair very minute ; at least a pair
of sessile papillae often present near tip of
tail. Spicules very dissimilar; short one
usually on the right, sword-shaped, 0.125
by 0.015 millimeter in size; left spicule
filiform, 0.740 millimeter long, of nearly
uniform thickness (0.006 millimeter)
throughout except at proximal end, where
it is dilated. Gubernaculum asymmetrical,
0.065 millimeter long.
Female 35.0 to 70.0 millimeters in length
by 0.20 to 0.35 millimeter in thickness at
middle of body. Posterior end behind
anus formed into a pointed, ventrally
curved tail (fig. 17, 6). Pharynx 0.058 to
0.072 millimeter long. Anterior portion
of oesophagus 0.46 to 0.78 millimeter long,
posterior portion 4.0 to 7.6 millimeters; total length of oeso-
phagus 4.5 to 8.4 millimeters or about one-eighth to one-ninth
of total body length. Distance from anterior end of worm to
cervical papillae 0.13 to 0.16 millimeter, to nerve ring 0.23 to
0.25 millimeter, to excretory pore 0.60 to 0.66 millimeter. Vulva
not prominent, behind middle of body. Distance from tip of tail
to vulva 2.6 to 5.9 millimeters, and to anus 0.17 to 0.21 milli-
meter. Vagina short, directed anteriorly from vulva and follow-
ed by long ovejector. Uteri divergent; anterior uterus becomes
receptaculum seminis near posterior end of oesophagus, the
posterior uterus being similarly modified behind level of vulva.
Ovaries much coiled. Eggs oval, embryonated at deposition, 54
G. 16. Gongylonema neo-
plasticum, anterior end,
lateral view.
578
The Philippine Journal of Science
1931
Fig. 17. Gongylonema neoplasticum. a, Posterior end of male, ventral view ; b, posterior end
of female, lateral view.
to 56 by 34 to 36 microns in size, with smooth shell about 3
microns in thickness.
Location. — Squamous-celled anterior portion of digestive tract,
usually beneath gastric mucosa.
Life history. — The development of this parasite requires an
intermediate host, which is invariably an insect. Cockroaches,
such as, Blatta orientalis, Phyllodromia germanica, and Peripla-
neta ameHcana; dung beetles, such as, Ateuchus, Aphodius, and
other genera of the family Scarabseidse ; cellar beetles and meal-
worm beetles of the family Tenebrionidse are all possible inter-
mediate hosts. Hall (1916) describes the life history as fol-
lows: The eggs of the worm are passed out of the body in
desquamations of the epithelium of the digestive tract with the
46,4 Tubangui: Worm Parasites of the Brown Rat 579
feces. If ingested by any one of the above insects, they hatch
in the intestine, and the liberated embryos, which measure 250
by 13 microns, follow a certain route and are finally found en-
capsulated in the musculature of the prothorax and legs of the
intermediary host. At this stage the larvae are 0.792 to 1.215
millimeters long and are coiled in spirals within their individual
cysts. They are rather slender and possess a conical tail that
often terminates in two or three papillalike projections of va-
riable size. Occasionally a wing-shaped prominence with
fringed or serrate edges is present. Anteriorly the larvae are
very similar in appearance to the mature worms, except that
the pharynx is relatively longer than in the adult and the oeso-
phagus is nearly as long as the intestine. In the beginning the
growth of the encapsulated larvse is faster towards the ante-
rior end, but later the rate of growth is reversed. The nerve
ring and excretory pore are distinct, the latter located halfway
between the former and the union of the two portions of the
oesophagus. Near the region where the vulva will later de-
velop in the female, the anlage of the reproductive system ap-
pears in the form of an oval body consisting of a number of cells
or a syncytium with several nuclei.
If an insect harboring these encysted larvae is ingested by a
proper vertebrate host, such as a rat, the latter are liberated
from their capsules due no doubt to the action of the gastric
juice, and on the following day they will be found to have
penetrated into the mucous membrane of the stomach and some-
times also into that of the oesophagus and tongue. During the
first ten days growth is rather slow, the larvae only doubling
their original length. They molt at about this time and their
tails become simple like those of the adult worms. Then they
grow more rapidly, and after two months the females begin to
deposit eggs.
References.— 4, 5, 19, 40, 43, 63.
Family RICTULARIIDJE Railliet, 1916
Subfamily RICTULARIIN^E Hall, 1913
Genus RICTULARIA Froelich, 1802
RICTULARIA WHARTONI sp. nov. Fig. 18.
This nematode is named in honor of the late Mr. Lawrence
D. Wharton, one of the early pioneers in the field of parasitology
in the Philippine Islands.
Description. — Male unknown.
580 The Philippine Journal of Science 1931
Female 25 to 33 millimeters in length by 0.65 to 0.90 millimeter
in thickness across middle of body. Cuticle transversely striated,
often swollen anteriorly forming a pair of ventrolateral cuti-
cular expansions 0.40 to 0.90 millimeter long (fig. 18, a).
Anterior end bent ventrally in preserved specimens, the rest of
body length turned towards opposite direction or rolled into a
semicircle; posterior end conical, ending in a short fine point
(fig. 18, c). Head 0.145 to 0.195 millimeter in thickness across
base of buccal capsule, provided with two ventral papillae. Buc-
cal capsule well developed, 0.05 to 0.07 by 0.06 to 0.08 milli-
meter in size, with its aperture surrounded by a series of den-
ticles (corona radiata) and its base armed with three conical
teeth possessing serrated borders (fig. 18, &). (Esophagus 3.5
to 4.6 millimeters long. Nerve ring 0.30 to 0.35 millimeter
from anterior end. Cervical papillae not very conspicuous, 0.70
to 0.74 millimeter from anterior end. There are 42 to 43 pairs
of "combs" extending from the head to the level of the vulva
and measuring 0.045 by 0.015 to 0.200 by 0.145 millimeter;
first pair of "combs" almost ridgelike, the rest bigger, more
distinct and gradually becoming more spinelike (fig. 18, a).
Behind the vulvar level there are 47 to 50 pairs of spines 0.05
to 0.16 millimeter long, the first three or five pairs being really
of a transitional type and the most posterior pair shorter; last
pair of spines immediately behind level of anal opening. Vulva
moderately prominent, usually in front (0.3 millimeter) of level
of posterior end of oesophagus, occasionally directly opposite or
even slightly behind this level. Vagina directed posteriorly from
vulva. Uteri convergent. Eggs with smooth fairly thick shell,
embryonated at time of deposition, measuring 44.2 to 47.5 by
34 microns. Anus 0.215 to 0.270 millimeter from tip of tail.
Specific diagnosis. — Rictularia: Male unknown. Female 25
to 33 millimeters in length by 0.65 to 0.90 millimeter in max-
imum thickness; with a pair of ventrolateral cuticular dila-
tations in cervical region. Base of buccal capsule armed with
three conical teeth possessing serrated borders. (Esophagus 3.5
to 4.6 millimeters long; distance from anterior end to nerve ring
0.30 to 0.35 millimeter; to cervical papillae 0.70 to 0.74 milli-
meter. Forty-two to forty-three pairs of "combs" from head
to level of vulva and forty-seven to fifty pairs of spines from
immediately behind vulvar level to posterior end of body. Vulva
usually in front of posterior end of oesophagus. Anus 0.215 to
0.270 millimeter from tip of tail. Eggs 44.2 to 47.5 by 34.0
microns in size.
46,4 Tubangui: Worm Parasites of the Brown Rat 581
K
m
Fig. 18. Rictularia whartoni sp. nov. a, Anterior end of female, lateral view; b, anterior
end of female, ventral view ; c, posterior end of female, lateral view.
Location. — Stomach and small intestine.
Locality. — Manila, P. I.
Type specimens. — Philippine Bureau of Science parasitologie-
al collection, No. 10.
Life history. — Unknown.
Reference. — 6, 19, 63.
582 The Philippine Journal of Science 1931
Class ACANTHOCEPHALA Rudolphi, 1808
Order ECHINORHYNCHATA Faust, 1929
Family MONILIFORMIDiE Van Cleave, 1924
Genus MONILIFORMIS Travassos, 1915
MONILIFORMIS MONILIFORMIS <Bremser, 1811) Travassos, 1915. Fig. 19.
Synonyms: Echinorhynchus moniliformis Bremser, 1811; Gigantor-
hynchus moniliformis (Bremser, 1811) Railliet, 1893; Hormorhyn-
chus moniliformis (Bremser, 1811) Ward, 1917; Echinorhynchus
cestodiformis Linstow, 1904.
This is appropriately known in ordinary language as the
beaded thorn-headed worm. In the adult stage it is a common
parasite of rats and other rodents and occasionally of dog and
man. Calandruccio (1888), who infected himself experimentally
by ingesting several infective larvae, was able to demonstrate
that the presence of the parasite in man in large numbers may
produce diarrhoea and severe gastrointestinal pain accompanied
by exhaustion, somnolence, and ringing of the ears. The expul-
sion of the worms with male fern caused the symptoms to dis-
appear two days after the treatment.
Description. — Body whitish or creamy-white in color, atten-
uated at both extremities, divided superficially except at ex-
treme anterior and posterior ends by annular grooves into a
series of beadlike pseudo-segments that give the worm a moni-
liform appearance. Size very variable in both sexes, the small-
est specimens usually immature. Proboscis (fig. 19, a) cylin-
drical, protrusible, relatively short, with broadly rounded distal
end; 0.425 to 0.670 by 0.15 to 0.21 millimeter in size, armed
with 12 to 16 longitudinal rows of recurved hooks, each row
composed of 7 to 12 hooks; hooks 24 to 30 microns long, each
with a single posteriorly directed root process (fig. 19, &).
Proboscis sheath large, 0.5 to 1.3 by 0.22 to 0.42 millimeters
in size, its wall composed of two muscular layers, of which the
outer is made up of diagonally wound fibers. Lemnisci filiform,
2.4 to 10.0 millimeters long, with few large nuclei.
Male 5.5 to 86.0 millimeters in length by 1.0 to 1.5 millimeters
in maximum breadth at middle of body ; posterior end expanded
into small bell-shaped bursa copulatrix, which, however, is
usually retracted within the body, being forced out only during
the copulatory act or as the result of the contraction of the wall
during the preservation of the specimen. Reproductive organs
46,4 Tubangui: Worm Parasites of the Brown Rat 583
0.05mm
a b c
FlG. 19. Moniliformis moniliformis, a, Anterior end, lateral view ; b, hooks ; c, egg.
at posterior portion of body cavity. Testes, of which there are
two, are oval, elongated, one immediately behind the other, 0.2
to 4.0 (usually 2.0) by 0.12 to 0.96 millimeters in size. Prostatic
glands eight in number, roundish to oval in shape, compressed
and crowded together behind testes, the entire mass measuring
in mature worms 0.45 to 3.60 by 0.25 to 1.10 millimeters.
Female 7 to 270 by 1.5 millimeters in size. Ovary present
only in larval stage, produces large numbers of ova which later
are found free in the body cavity of the adult worm. Eggs
ellipsoidal, 109 to 137 by 40 to 63 microns in size, and provided
with three envelopes; in fully mature eggs outer shell slightly
wrinkled and the inclosed embryo brown or dark-colored,
striated and covered with minute spines (fig. 19, c).
Location. — Small intestine.
Life history. — Indirect, the intermediate hosts being species
of beetles (Blaps mucronata), cockroaches (Periplaneta ante-
ricana) , and possibly other insects. If ingested by these insects
the eggs develop into oval larvse in their abdominal cavities.
Each larva is inclosed in a very delicate cyst, which, according
to Southwell (1922) is easily lost. The larva on being swal-
lowed by a suitable mammalian host together with the insect har-
584 The Philippine Journal of Science 1931
boring it escapes from its cyst (if this has not already been lost)
and develops directly into an adult worm. The mode of in-
fection in man is somewhat obscure; it may result from the ac-
cidental ingestion of either of the infected intermediate host
or food polluted by cysts from disintegrated cockroaches and
beetles.
Prevention. — Consists in the destruction of rats and mice
that play the role of reservoirs and of cockroaches and beetles
that act as intermediate hosts of the parasite. Foods should be
protected from these insects.
References.— 14, 46, 47, 51, 53, 54, 56.
SUMMARY
Besides the role that they play as carriers and reservoirs of
bubonic plague and other bacterial as well as spirochetal in-
fections, rats often harbor parasitic worms, some of which are
also a menace to human health. In view of this and because
of the fact that the helminthic fauna of rats has never been
studied extensively in the Philippine Islands, it seemed desir-
able to undertake a systematic examination of these animals
for the purpose of finding out if they are infested with parasites
that are transmissible to man.
The examination of nine hundred fifty rats (Mus norvegicus)
resulted in the identification of the following sixteen species of
helminths: Trematodes: Euparyphium ilocanum, E. guerreroi,
and E. murinum sp. no v.; cestodes: T tenia tseniaformis (larval
form), Raillietina garrisoni sp. nov., Hymenolepis diminuta, and
H. nana; nematodes: Gongylonema neoplasticum, Hepaticola
hepatica, Heterakis spumosa, Nippostrongylus muris, Protospi-
rura muricola, Rictularia whartoni sp. nov., Strongyloides ratti
and Trichosomoides crassicauda; Acanthocephala: Moniliformis
moniliformis.
The following parasites of rats have been reported from hu-
man beings : Euparyphium ilocanum, Hymenolepis diminuta, H.
nana, Syphacia obvelata, Hepaticola hepatica, and Moniliformis
moniliformis. The first four species mentioned in this para-
graph have been reported to occur in man in the Philippine
Islands.
It is also believed that Raillietina garrisoni should be included
among the parasites of the rat that are transmissible to man
because of its common occurrence and its close morphological
*'4 Tubangui: Worm Parasites of the Brown Rat 585
resemblance to the human tapeworm described by Garrison in
1911 from the Philippines as Davainea madagascariensis.
The morphology and the life history, if known, of each of the
different parasites are given and, in the case of the forms that
are transmissible to man, methods of avoiding infestation are
discussed.
ADDENDUM
After the manuscript of the above paper was submitted for
publication, I found in the literature a description by Hoeppli 2
of a new nematode, Rictularia tani, from the brown rat in Amoy,
China, with which Rictularia whartoni Tubangui should be com-
pared. The two forms resemble each other in several import-
ant characters, such as, in the number of their cuticular combs
and spines, the length of the oesophagus, and the location of
the nerve ring, vulva and anus. They differ in the presence of
a pair of ventrolateral cuticular dilatations in R. whartoni and
in the fact that the last pair of spines of R. whartoni is found
behind the anus, that of R. tani occurring in front of that level.
Because of these differences it is decided to maintain the Phil-
ippine Rictularia as a separate species.
Very recently there also came to hand a paper by Lopez-
Neyra3 that has an important bearing on the discussion of
Raillietina garrisoni. I described this as a new species of rat
tapeworm for, while recognizing its close alliance to Raillietina
celebensis (Janicki) Meggitt and Subramanian, 1927, it differs
from the latter in the number of its testes and uterine egg cap-
sules and in the size of its cirrus pouch. I also gave reasons
for suspecting its possible identity with Garrison's Davainea
madagascariensis which, according to Joyeux and Baer, differs
from the specimens described under the same name by other
observers. Now, according to Lopez-Neyra, the following rep-
resent one and the same species of parasite that should be known
as Kotlania madagascariensis (Davaine, 1869) : the collections
in the Parasitological Laboratory of the University of Paris
denominated as Type No. 108 (Davaine), No. 109 (Davaine),
No. 8 (Blanchard, Port-Louis) and No. 33 (Nossi-Be, 1873) ;
Taenia madagascariensis Leuckart, 1891 ; Davainea madagasca-
riensis Garrison, 1911; D. formosana Akashi, 1916; Raillietina
•Centralbl. 1 Bakteriol. u. Parasitenk. 1 Abt Orig. 110 (1929) 75-78.
8 Ann. Parasit. Hum. et Comp. 9 (1931) 162-184.
264209 4
586 The Philippine Journal of Science 1931
celebensis (Janicki) Meggitt and Subramanian, 1927; R. fune-
bris Meggitt and Subramanian, 1927 ; and possibly R. fluxa Meg-
gitt and Subramanian 1927. If Lopez-Neyra's hypothesis is
accepted, then Raillietina garrisoni will have to fall in line with
the above synonymy.
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ILLUSTRATIONS
[Drawn by V. V. Marasigan under the direction of the author.]
TEXT FIGURES
Fig. 1. Euparyphium ilocanum. a, Entire worm, ventral view; 6, ante-
rior end, showing arrangement of spines on cephalic collar,
ventral view. (After Tubangui, 1931.)
2. Euparyphium guerreroi. a, Entire worm, ventral view; b, ante-
rior end, showing arrangement of spines on cephalic collar,
ventral view. (After Tubangui, 1931.)
3. Euparyphium murinum sp. nov. a, Entire worm, ventral view;
6, anterior end, showing arrangement of spines on cephalic
collar, ventral view.
4. Taenia tseniuformis. a, Entire larva (after Sambon, 1924) ; 6,
scolex, anterior view; c, rostellar hooks.
5. Raillietina garrisoni sp. nov. a, Rostellar hooks; 6, scolex; c,
mature segment; d, gravid segment; e, egg.
6. Hymenolepis diminuta. a, Head; 6, mature segment, dorsal view;
c, gravid segment; d, egg.
7. Hymenolepis nana, a, Entire worm (from Ransom, 1904) ; 6, ros-
tellar hooks.
8. Hymenolepis nana, a, Head; 6, mature segment, ventral view;
c, gravid segment; d, egg.
9. Strongyloses ratti, entire worm, a, Anus, oe, oesophagus; ovf ova-
ry; v, vulva.
10. Trichosomoides crassicauda. a, Mature female with male in ute-
rus (after Hall, 1916), m, male worm; v, vulva;, 6, anterior
end of mature male (after Thomas, 1924) ; c, egg.
11. Hepaticola hepatica. a, Anterior end of mature female (after
Nishigori, from Yorke and Maplestone, 1926); oe, oesophagus;
v, vulva; 6, egg.
12. Nippostrongylus muris. a, Anterior end, lateral view; b, bursa,
dorsal view.
13. Syphacia obvelata. a, Anterior end of female, lateral view; 6,
male, lateral view; c, posterior end of male, ventral view; df
posterior end of female, lateral view; e, egg. (All from Yorke
and Maplestone, 1926.)
14. Heterakis spumosa. a, Anterior end of female, ventral view;
by posterior end of female, lateral view; c, posterior end of male,
ventral view.
15. Protospirura muricola. a, Anterior end of female, ventral view;
b, mouth, anterior view; c, posterior end of male, lateral view;
d, posterior end of female, lateral view.
16. Gongylonema neoplasticum, anterior end, lateral view.
17. Gongylonema neoplasticum. a, Posterior end of male, ventral
view; 6, posterior end of female, lateral view.
18. Rictularia whartoni sp. nov. a, Anterior end of female, lateral
view; 6, anterior end of female, ventral view; c, posterior end
of female, lateral view.
19. Moniliformis moniliformis, a, Anterior end, lateral view; 6,
hooks; c, egg.
591
NOTES ON DENGUE
By R. L. Holt and J. H. Kintner
Of the United States Army Medical Department Research Board
Bureau of Science, Manila
TWO TEXT FIGURES
The question of immunity in dengue has been discussed on
innumerable occasions and much research work has been done
on this phase of the disease. That immunity to the disease
does occur was reported by Simmons, St. John, and Rey-
nolds,1 they reporting an effective resistance to reinfection in
the case of thirty-six volunteers who had previously suffered
from experimentally induced attacks of dengue. The immunity
was proven in these cases at periods from one-half to thirteen
months after the attack. This finding has been supported by
experimental work carried on by the authors in six cases, all
proving immune to the same strain with which they were origi-
nally infected twenty days previously. Following the proof of
this immunity an attempt was made to reinfect these volun-
teers by feeding on them mosquitoes that had been infected by
feeding on two other cases of dengue fourteen days previously.
Reinfection was not accomplished in any of the six cases al-
though they were all held twenty-one days after being bitten
by the mosquitoes infected from an outside source.
This attempt to reinfect from outside sources was suggested
by two cases that came under our observation and that, coupled
with the fact that so many cases are immune to the virus with
which they were originally infected, led us to suspect that there
might possibly be different strains of dengue virus. We still
believe in that possibility and attempts are being made to secure
dengue-infected mosquitoes from other localities to prove or
disprove this possibility.
The two cases mentioned above have the following histories :
1. This case was an officer's wife who arrived in the Phil-
ippine Islands in March, 1930, and has lived in Manila since
Philip. Journ. Sci. 44 (1931) 174.
593
594 The Philippine Journal of Science i»3i
that time. She developed dengue in March, 1930, July, 1930,
and October, 1930. All three attacks were typical and severe.
There was a marked reduction in the leucocyte count, a crossing
of the staff and segmented leucocytic curves and a temperature
curve characteristic of the disease. In spite of the fact that
transmission experiments were not carried out, all other find-
ings strongly supported the belief that in this case three attacks
of dengue occurred in less than seven months.
2. This case was a volunteer who was kept in a screened cubi-
cle in Sternberg General Hospital for eight days, preliminary ob-
servation, during which time no symptoms of illness appeared.
On the ninth day this volunteer was bitten by fifty mosquitoes
selected at random from a cage of five hundred insects that had
been infected from an experimentally induced case of dengue
eighteen days previously. Three other volunteers were infected
at the same time by the bites of other insects from the same lot
and all three developed typical dengue. The case in discus-
sion did not develop dengue after fourteen days, and forty-six
mosquitoes of another infected lot were allowed to bite him.
During three weeks observation following the second feeding no
dengue occurred and he was discharged from hospital as im-
mune. During his stay in hospital he had been transferred to
a station 30 miles outside Manila and joined that station imme-
diately after leaving hospital. Fourteen days after joining his
new station he developed a typical case of dengue, as was shown
by blood findings and clinical symptoms. There was no ques-
tion as to the infectivity of the two lots of mosquitoes used in
this case, and it is not believed that infection could have been
produced as a result of feeding upon him the last lot of mosqui-
toes thirty-five days previously. No such prolonged incubation
period has been observed, the vast majority of cases developing
the disease within eight days after infection. There are only
three possible conclusions in this case: (1) The extremely re-
mote contingency that there was a long delayed incubation pe-
riod after the second infection by mosquitoes; (2) that he lost
a high degree of immunity in about twenty-nine days (allow-
ing six days incubation period for the development of the dis-
ease) ; (3) that there are different strains of dengue virus.
Our experience and the experience of others lead us to believe
that the first conclusion is practically untenable and that the
second is not probable.
The first case related has no parallel insofar as we have been
able to find, this patient having had three proven attacks of
46, 4 Holt and Kintner: Notes on Dengue 595
dengue, severe in character, in a period of seven months. The
evidence in these two cases certainly leads us to suspect the
possibility of strains of dengue virus. Other cases not so strik-
ing as are these two tend to confirm our belief.
St. John 2 has devised a feeding cell so constructed that normal
mosquitoes may be fed on other mosquitoes ground up and sus-
pended in blood. In an attempt to attenuate the virus of dengue
we have made serial feedings of normal Aedes segypti on macer-
ated infected Aedes, using fifty infected mosquitoes ground
up in one cubic centimeter of nonimmune blood. These Aedes
infected by feeding were allowed an incubation period of four-
teen days when they in turn were fed to normal Aedes in the
same manner as above. Five transfers were made and in all
cases the insects were well filled with the blood mixture after
feeding. Fourteen days after the third transfer dengue was
produced in a susceptible volunteer, showing that transfer from
mosquito to mosquito can be accomplished at least three times
without the intervention of a human host. It is unlikely, of
course, that this occurs in nature. The dengue produced by the
bites of mosquitoes that had been infected by the third serial
transfer from insect to insect was not modified insofar as we
could determine, there being a reduction of the total white count
to five thousand per cubic millimeter, a maximum temperature of
104° F., and a typical crossing of the segmented and staff forms
of leucocytes. The subjective symptoms were as severe as in
the ordinary attack of dengue. Fourteen days after the fifth
serial transfer one hundred forty-five mosquitoes of this lot were
allowed to bite a susceptible volunteer and did not produce
dengue in twenty days. Subsequently this volunteer was bitten
by a known infected lot of mosquitoes and developed typical
dengue, showing that the volunteer was not immune and that
the dengue virus had been lost in the five direct transfers from
mosquito to mosquito or that it had been attenuated to such an
extent that infection did not occur.
In another series of experiments attempts to attenuate the
virus were made as follows :
Two volunteers were subjected to a preliminary observation
period of one month in a screened cubicle. Dengue was pro-
duced experimentally in a third volunteer and on the first day
of the fever 10 cubic centimeters of blood was removed, allowed
to clot, the serum separated and immediately frozen at a tem-
2 Op. cit, Plate 2.
596
The Philippine Journal of Science
1931
perature of — 7° F. Twenty-four hours later 0.3 cubic centi-
meter of this serum was injected subcutaneously into the first
volunteer and forty-eight hours after freezing 0.3 cubic centi-
meter of the same serum was injected into the second volunteer.
The results obtained in the volunteer who received 0.3 cubic
centimeter of the serum that had been frozen twenty-four hours
were inconclusive in that the incubation period was too short to
be entirely certain of the source of the infection, but in the
volunteer receiving the virus frozen for forty-eight hours the
disease developed on the seventh day and was typical, as shown
by fig. 1.
Fig. 1. Experimental dengue in American volunteer 28.
46,4
Holt and Kintner: Notes on Dengue
597
From this experiment it seems certain that the virus of den-
gue is affected little, if at all, by freezing at — 7° F. for a period
of forty-eight hours.
A number of investigators have attempted to modify the
course of dengue by the use of "convalescent" serum and all
report failures. We have gone a step further. From each of
four volunteers who had recovered from experimental dengue
10 cubic centimeters of plasma was removed on the fourteenth
day after fever had disappeared. The pseudoglobulins were
precipitated out of the pooled plasma, diluted with 0.85 per cent
salt solution, and injected into a volunteer on the first day of
dengue. No modification of the disease was noted. As shown
by the chart, fig. 2, there was a primary and secondary rise of
temperature, the height of the first reaching 103° F. and the
EXPERIMENTAL DENGUE A.V. No. 8© COX f CLARE MCE T
'
FIRST EXPERIMENT.
JAW. 15 - FED 63 A. AEGYPTI
OF LOT 35-
ARROW SHOWS INJECTION OF PSEUDOGLOBULINS
SUSCEPTIBILITY TE3T. JAN. 26-
FED 92 INFECTED A. AEGYPTI OF LOT 37.
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598 The Philippine Journal of Science
second 101° F. There was no discoverable abatement of the
subjective symptoms, and none of the other symptoms seemed
to be modified.
CONCLUSIONS
There is certainly an immunity to dengue fourteen days after
an attack, both to the original infecting medium and to the virus
of two outside cases.
Peculiarities in the behaviour of attacks of dengue, two cases
of which are reported, coupled with the fact that immunity to
the homologous strain has been proven numbers of times lead
us to believe in the existence of "strains" of dengue virus.
The virus of dengue was tranferred from mosquito to mos-
quito by feeding for three transfers but was lost before the fifth
transfer.
There was no discoverable attenuation of the virus as a result
of the three transfers above mentioned.
Freezing dengue serum at — 7° F. for forty-eight hours did
not destroy the virus nor was it attenuated insofar as we could
discover.
Concentrated "immune bodies" from the serum of recovered
cases of dengue do not affect the course of the disease when
injected on the first day of the attack.
ILLUSTRATIONS
TEXT FIGURES
Fig. 1. Experimental dengue in American volunteer 28.
2. Experimental dengue in American volunteer 26.
599
RESISTANCE OF DENGUE VIRUS
By R. L. Holt, Wm. D. Fleming, and J. H. Kintner
Of the United States Army Medical Department Research Board
Bureau of Science, Manila
FOUR TEXT FIGURES
Considering the enormous amount of work that has been done
on dengue, it is rather surprising to note that comparatively
little has been done on the resistance of dengue virus to outside
influences. Cleland, Bradley, and McDonald x were able to pro-
duce dengue by inoculating infective blood which had been stored
at refrigerator or room temperature for periods up to 172 hours.
Blanc and Caminopetros 2 found that the addition of relatively
small amounts of bile killed the virus in a short time. The virus
has been preserved for periods of several days by different meth-
ods but almost always at low temperature. Drying of the virus
seems to destroy it much more quickly than when it is kept moist
and subjected to the same procedures. A temperature of 55° C.
will render the virus noninfective within a period well under
thirty minutes. It is certain that the virus resists the actions
of the body fluids for an incubation period of six to nine or ten
days and for at least three days after the onset of the disease.
The mixing of convalescent sera and virus in serum does not
seem to affect the infective power of the virus. It is well estab-
lished that the virus lasts throughout the life of the infected
Aedes. We have demonstrated that the virus does not remain
infective at the end of seven days at 37.5° C. in Tyrode's medium
alone or with the addition of fresh testicular tissue from a rabbit.
At just what period it lost its infectivity we are unable to say.
We have already reported that freezing dengue serum at ■— 7° C.
for a period of forty-eight hours does not seem to affect its in-
fective power in the least.
Since it is well known that ultra-violet rays will destroy bac-
teria of many kinds in water and that X-rays will destroy the
spores of many of the fungi, it was decided that it was worth
1 Journ. Hyg. Cambridge 18: 217.
2 Bull. Acad. Med. Athens 26: 37.
264209 — 5 601
602
The Philippine Journal of Science
1931
while to see if either of these agents would affect the virus of
dengue in the body of the mosquito. As a preliminary step it
was necessary to determine the resistance of Aedes zegypti to
the effects of X-ray and ultra-violet light. The insects were
placed in a glass feeding cell, the open end of which was covered
with mosquito netting to allow passage of the rays. In all ex-
posures the number of insects used was twenty-five. In X-ray
exposures the setting of the machine was constant; namely, 70
K.V., 5 M.A. and the distance from the center of the feeding cell
was 30 centimeters. Only the time varied. Almost all of the
mortality among the insects exposed occurred within a few min-
utes of the exposure and the mortality was not even roughly
proportional to the length of exposure. For instance, when
exposed to approximately one-twelfth erythema dose (30 seconds)
four insects were dead in fifteen minutes and only two more
had died at the end of twenty-four hours, one-sixth erythema
dose (60 seconds) showed five dead in fifteen minutes and only
two more dead in twenty-four hours, one-third erythema dose
(90 seconds) showed eight and eleven dead, while one-half ery-
thema dose (3 minutes) showed figures of two and naught. In
this manner it was found that these insects could readily stand
as much as two erythema doses with about 50 per cent living
after twenty-four hours. A few lived for several days after
this exposure.
Table 1. — Aedes segypti exposed to X-ray,
Setting.
Number
exposed.
Time.
Dis-
tance.
Living at end
of—
15 min-
utes.
24 hours.
70 KV~5 MA _
25
25
25
25
sec.
30
60
90
180
cm.
30
30
30
30
21
20
17
23
19
18
14
23
70KV-5MA.
70 KV-5 MA
70KV-5MA _ _
In the same manner experiments were carried out to show
the effects of ultra-violet on the normal Aedes. The source of
the ultra-violet was a Cooper Hewitt unit. The container for
the mosquitoes in all experiments was a round glass feeding cell
14 centimeters deep with the open end toward the source of light
covered by mosquito netting. In all cases there were 65 volts
46,4 Holt et al.: Resistance of Dengue Virus 603
across the arc of the mercury lamp and the current was 3.5
amperes.
This arc gave light of a total energy at 40 centimeters of 22
microwatts per square millimeter, or 0.000005 gram calories per
second per square millimeter.
The light had the following spectral characteristics :
_,.„. . Microwatts
Millimicrons. Per cent. per square
millimeter.
185-1400 100 22
250-1400 78 17
310r1400 66 15
185-250 22' 5
185-310 34 8
250-310 12 3
With an exposure of fifteen minutes at a distance of 40 centi-
meters, thirty-four insects showed no ill effects in seventy-two
hours.
With thirty minutes exposure at 25 centimeters distance, of
twenty-five mosquitoes two were dead at the end of the exposure
and seven more were dead at the expiration of thirty minutes
after the exposure. These deaths may have been due to heat
or ultra-violet or both.
Of thirty-four mosquitoes exposed for one hour at a distance
of 18 centimeters, with the light filtered through 4 millimeters
of quartz and 1 centimeter of water and screened from the heat
of the lamp, nine were dead in twenty-four hours and twenty-
five were living but very weak. At the end of forty-eight hours
thirty-three were dead and one was living but very weak.
With a forty-five minute exposure under the same conditions,
of thirty mosquitoes two were living and twenty-eight dead at
the end of twenty-four hours, and at the end of forty-eight hours
all were dead.
Of twenty-five mosquitoes exposed for thirty minutes under
the same conditions as related in the preceding experiment, four-
teen were living at the end of twenty-four hours, three were
living at the end of forty-eight hours, one was living at the end
of seventy-two hours, and all were dead at the end of ninety-
six hours. The fourteen living at the end of twenty-four hours
were so weak that they were unable to feed on an animal.
To the ultra-violet that had passed through 4 millimeters of
quartz, 1 centimeter of water and 2.5 millimeters of window
glass which cut off at about 310 millimicrons, thirty Aedes were
604
The Philippine Journal of Science
1931
exposed for forty-five minutes. All were living after twenty-
four hours, and eighteen of the thirty fed on a guinea pig. Five
were dead at the end of forty-eight hours. No others died until
after one hundred fourteen hours when twenty-four of the thirty
were living and at the end of one hundred sixty-eight hours
twenty-three were living.
Thirty Aedes were exposed for fifteen minutes with the light
filtered through 4 millimeters of quartz and 1 centimeter of
water. After twenty-four hours, twenty-seven were living and
nineteen fed on a guinea pig. No more died until the end of one
hundred twenty hours, when a total of four were dead. None
died between one hundred twenty and one hundred forty-four
hours.
Table 2. — Mosquitoes exposed to ultra-violet light.
[Voltage 65 and current 3.5 amperes in all the following cases.]
Dis-
tance.
Time.
Filter.
Num-
ber ex-
posed.
Living at end of —
m
o
c
g
©
CO
w
o
X\
O
Xi
00
3
o
X
C<l
o
CO
Oi
2
O
■*#
T-t
o
A
00
to
cm.
40____
25__._
18„_.
18___.
18_
18_._ _
18____
min.
15
30
60
45
30
45
14
0
0
4 mm quartz
a n d 1 cm
water.
4 mm quartz
a n d 1 cm
water.
4 mm quartz
and 1 cm
water.
4 mm quartz,
1 cm water,
and 2.5 cm
window
glass.
4 mm quartz
a n d 1 cm
water.
34
25
34
30
25
30
30
34
23
16
25
2
14
30
27
1
0
3
25
1
0
24
26
23
From these experiments it was thought that the maximum
dose of X-ray for mosquitoes was two erythema doses if the
insects were to be able to feed on a volunteer after exposure
46'4 Holt et aL: Resistance of Dengue Virus 605
and that the maximum exposure to ultra-violet would be about
fifteen minutes.
Four volunteers, A.V. 34-Immordino, A.V. 35-Hawley, A.V.
36-Cook, and A.V. 37-Small were admitted to screened cubicles
in Sternberg General Hospital February 27, 1931, and submitted
to an observation period of thirteen days, during which time
they showed no evidence of illness. All were recent arrivals in
the Philippine Islands and gave no history of dengue. At 8 a.
m. on the fourteenth day four lots of proven infective mosquitoes
were treated as follows:
Lot 1 consisted of forty-six mosquitoes. They were exposed
for fifteen minutes to ultra-violet produced by a mercury arc
lamp with 65 volts crossing the arc and showing a current of
3.5 amperes and with a distance of 40 centimeters from the
front of the feeding cell, the light being filtered through a cell
of 4 millimeters of quartz and 1 centimeter of water. This cell
passed light from 185 to 1,400 millimicrons wave length. Most
of the insects congregated at the back of the feeding cell during
the exposure so that the distance was about 50 centimeters from
the arc to where the majority of the insects congregated. Many
of the insects fell to the bottom of the feeding cell during the
exposure but ten minutes later all had apparently recovered.
One hour later three had died, seventeen refused to feed, and
twenty-six fed on A.V. 34-Immordino. Six days later this vol-
unteer developed typical dengue as shown by the chart (fig. 1).
Of this lot of mosquitoes sixteen survived for nine days.
Lot 2 consisted of forty-six infected Aedes and were subjected
to the same conditions as outlined in lot 1, except that a window-
glass filter, passing light from 310 to 1,400 millimicrons wave
length, was inserted between the source of ultra-violet and the
insects. Exposure time was fifteen minutes and distance 40
centimeters. One hour later two were dead, eighteen refused to
feed, and twenty-six fed on A.V 36-Cook. Five days later this
volunteer developed dengue as shown by the chart (fig. 2).
Twenty-seven of these insects survived for nine days.
Lot 3 consisted of forty-six infected Aedes that were treated
as follows: An X-ray machine was set at 70 K.V., 5 M.A. and
with a distance of 30 centimeters. The insects were radiated for
six minutes with this setting, which exposed them to approxi-
mately one erythema dose. One hour later seven were dead,
fifteen refused to feed, and twenty-four fed on A.V. 35-Hawley,
606
The Philippine Journal of Science
1931
Fig. 1.
Chart of American volunteer 34-
Immordino.
Fig. 2.
Chart of American volunteer 36-
Cook.
who developed dengue six days later, as shown by the chart
(fig. 3). Seven mosquitoes survived nine days.
Lot 4 consisted of forty-two infected Aedes that were subjected
to X-ray radiation equal to approximately two erythema doses,
the setting of the machine being the same as for lot 3 and the
time doubled. At the end of one hour six were dead, seven re-
fused to feed, and twenty-nine fed on A.V. 37-Small, who devel-
oped dengue on the sixth day, as shown in the chart (fig. 4).
Only one mosquito survived nine days.
The insects showed great perturbation while being treated with
X-ray and ultra-violet unscreened, but were not visibly dis-
turbed by treatment with ultra-violet when a window-glass
screen was interposed.
The disease produced in the above-mentioned volunteers was
not modified insofar as we could determine.
46,4
Holt et al.: Resistance of Dengue Virus
607
iNFfcUTlVH Y IE3T. riAKCrt 13-
FED 24 INFECTED A.AEGYPT1 OF LOT 51.
WFECTI0U3 MATERIAL. MARCH I3-PED 29
INFECTED A.AEGYPT1 OF LOT 51
1NCUBA7
'ION 1 DENGUE FEVER
IMCUBAT
ON. 1 DENGUE FEVER,
2 105
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Hawley.
Chart of American volunteer 37-
Smali.
CONCLUSIONS
Aedes segypti shows much greater resistance to the effects of
ultra-violet light and X-ray than does man.
Dengue virus does not appear to be adversely affected by rela-
tively large amounts of X-ray and ultra-violet, despite the fact
that it is well known that several species of bacteria and molds
are affected by ultra-violet light and that several of the fungi
and their spores are affected by X-ray.
It seems apparent that no good can be hoped for in the treat-
ment of dengue cases by the use of either of the above-mentioned
forms of energy.
ILLUSTRATIONS
TEXT FIGURES
Fig. 1. Chart of American volunteer 34-Immordino.
2. Chart of American volunteer 36-Cook.
3. Chart of American volunteer 35-Hawley.
4. Chart of American volunteer 37-Small.
609
THE ATTEMPTED CULTIVATION OF MYCOBACTERIUM
LEPRAE
By Wade W. Oliver
Visiting Professor and Head, Department of Sanitary Bacteriology and
Immunology, School of Hygiene and Public Health
University of the Philippines
Walprido de Leon
Professor of Pathology and Bacteriology, College of Medicine; Professorial
Lecturer on Immunology, School of Hygiene and Public
Health, University of the Philippines
and
Alfredo Pio de Roda
Instructor in Hygiene, College of Medicine, University of the Philippines
one plate
The immediate activator of the present studies was the re-
ported cultivation of Mycobacterium leprse in 1929 by Wherry 1
in Manila and by Shiga2 in Chosen, Korea.
In Wherry's studies, made in the School of Hygiene and Public
Health in Manila, he reported microscopic proliferation of M.
leprse in cultures from three cases. One loopful of blood, ob-
tained by the routine "snip" method, was inoculated into the
water of syneresis of slants of glycerinized ovomucoid yolk
agar, to which had been added 1 to 2 drops of autoclaved oleic
acid and 1 to 2 drops of autoclaved 10 per cent dextrose solu-
tion in distilled water. Wherry reports that the best growth
was obtained in cultures that were kept for a month at 35 to
37° C. at partial oxygen tension (diminished 02 and increased
C02), a condition that was brought about by attaching the cul-
ture tubes by means of rubber tubing to agar slants inoculated
with B. coli. After one month at partial oxygen tension, a fine
syringe needle was inserted through the connecting rubber tub-
ing and the point of the needle buried in the cotton plug of
1 Journ. Inf. Dis. 46 (1930) 263.
'Journ. Chosen Med. Assoc. 19 (1929) 1.
611
612 The Philippine Journal of Science 1931
the culture tube, thereby establishing an 02 and C02 environ-
ment.
Shiga used, as the source of his inoculum, excised leprous
nodules that were ground in a 5 per cent solution of sulphuric
acid. The suspension was next incubated for twenty minutes,
and then centrifuged; the sediment was used for inoculation.
The medium employed by Shiga consisted of 4 per cent glycerin
bouillon potato wedges placed in Roux test tubes (reaction pH
6.8 to 7.0) with the lower portion of the tube, below the con-
striction, filled with 4 per cent glycerin bouillon. Shiga reported
growth of M. leprse after two months of aerobic incubation at
37° C, but says : "Die Kolonien auf Kartoffeln sind zart, dtinn
und unsichtbar." However, he says that upon transplant to
glycerin agar small, but visible, colonies developed.
The total number of cases cultured by us was twelve ; all were
active, recently admitted cases of leprosy at San Lazaro Hos-
pital, Manila, that had received no treatment, with the exception
of cases 11 and 12. We wish here to record our appreciation
of the fine spirit of cooperation manifested by Doctor Velasco
in placing these cases at our disposal. In the appendix will be
found a brief description of each case, and the direct smear
findings.
In all of the cases, the method of obtaining culture material
was the same. The skin over the leprous lesion was treated
with iodine, followed by alcohol ; then the area to be incised was
grasped with sterile forceps, and the incision made with a
sharp sterile knife. Next, the interior was scraped with the
tip of a second sterile knife, and blood containing lepra bacilli
was obtained on the edge of the knife. In many instances,
pieces of tissue were also obtained. By means of a sterile pla-
tinum loop a loopf ul of blood or a piece of tissue was transferred
immediately to the water of syneresis in the culture tube; in-
oculations on the surface of the slant, just above the water of
syneresis, were made as well. In each case, direct smears were
made from blood or tissue scrapings approximately equal in
amount to that employed as the inoculum of a single culture
tube. Direct smears, as well as smears from cultures, were
stained with steaming, and with unwarmed, carbol-fuchsin, fol-
lowed by acid alcohol and a counter stain of dilute LofHer's
methylene blue.
The media employed in our cultural studies were as follows :
(1) 4 per cent, 5 per cent, and 6 per cent glycerin ovomucoid
yolk agar containing oleic acid and dextrose (Wherry medium) ;
46'4 Oliver et al: Mycobacterium leprse 613
(2) 4 per cent glycerin bouillon Irish potato wedges (Shiga
medium) ; (3) 4 per cent glycerin bouillon lakatan (banana)
wedges; (4) 4 per cent glycerin bouillon saba (banana) wedges;
(5) 4 per cent glycerin bouillon sweet potato wedges; (6) 4
per cent glycerin bouillon gabi (tuber) wedges; (7) 4 per cent
glycerin bouillon ubi (tuber) wedges; (8) lakatan agar slants
(pH 7.2), made by mixing one part of ground 4 per cent gly-
cerin lakatan with two parts of 2.5 per cent meat infusion
agar; (9) serum glucose cystine agar slants and columns (Fran-
cis),3 to which in certain series 1 drop of normal human serum
was added to the water of syneresis, and to others 1 drop of sy-
philitic serum; and (10) infant human brain agar slants, made
by mixing one part of ground 4 per cent glycerin brain with
one part of 3 per cent nutrient agar, plus 2 drops of 10 per
cent dextrose solution and 2 drops of nucleic acid to each 3 cubic
centimeters of medium. In addition, slants of plain glycerin
and blood agar, as well as of Loffler's blood serum, were employed
as controls.
All of our cultures, primary cultures as well as transplants,
were incubated continuously at 37° C. for periods ranging from
eleven weeks to seven months, with the exception of one series
that was kept at room temperature. In all instances, parallel
culture tubes were incubated (a) aerobically, uncapped and
covered with a rubber cap; (b) anaerobically, by means of pyro-
gallic acid and potassium hydroxide; (c) partial oxygen tension
and increased C02. In certain of the partial oxygen tension
series, the original B. subtilis or B. coli was allowed to remain
unchanged during the entire period of incubation; in others,
freshly inoculated slants were attached to the culture tubes
every twenty-four hours. In other series, after four to six
weeks incubation under diminished oxygen and increased C02,
oxygen was admitted by inserting a fine needle through the con-
necting rubber tubing, and burying the point in the cotton
stopper of the culture tube.
A detailed analysis of the cultural findings in each case that
we studied will not be attempted. Rather, we will endeavor
briefly to review our results as a whole, with illustrative ref-
erences to certain typical cases.
Aerobic cultures, both capped and uncapped, have shown a
shorter persistence of M. leprse, in smaller numbers, than have
3 Public Health Reports 38 (1923) 1396.
614 The Philippine Journal of Science 1931
the comparable partial tension and anaerobic cultures. Aerobic
cultures on glycerin bouillon lakatan wedges, glycerin bouillon
saba, glycerin bouillon sweet potato, glycerin bouillon gabi, gly-
cerin bouillon ubi, lakatan agar slants, serum glucose cystine agar
slants, as well as slants of plain glycerin and blood agar and
Loffler's blood serum, have revealed a quite regular disappear-
ance of the implanted acid fasts within two weeks of incubation
at 37° C, and in a shorter time at room temperature. There is
a relatively brief persistence of the microorganisms with typical
staining reaction, followed by a rather rapid loss of acid-fastness,
and then by autolysis. Somewhat irregular persistence of M.
lepras for a longer period of time was observed in certain cases
in aerobic 37° C. cultures on infant human brain agar, glycerin
ovomucoid yolk agar (Wherry medium), and glycerin bouillon
Irish potato wedges ; in cases 1, 4, 5, 6, 7, 8, 9, and 10, the orig-
inal aerobic cultures revealed no acid fasts after one month at
37° C. In case 2 (Claudio Quinto), in which direct smears
from the lesion showed enormous numbers of M. leprse, a smear
from a fragment of blood and tissue scraping planted on an
aerobic slant of infant human brain agar revealed brightly stain-
ing, typical acid fasts in pure culture at the end of forty-eight
days' incubation at 37° C. The acid fasts were not, however, in
as large numbers as in the comparable partial tension culture,
and they disappeared on longer incubation. In the same case 2,
a comparable smear from the same amount of inoculum on aero-
bic glycerin bouillon Irish potato showed somewhat smaller num-
bers of equally typical acid fasts in pure culture upon forty-three
days' incubation at 37° C, which disappeared on further incu-
bation; whereas a comparable smear from the same amount of
inoculum in the water of syneresis and on the surface of a slant
of glycerin ovomucoid yolk agar (Wherry medium) revealed no
acid fasts at the end of forty-three days' incubation at 37° C.
On the other hand, in case 3 (Pablo Carpio), which likewise
showed M. leprse in enormous numbers in direct smears from
the lesion, a smear from the glycerin bouillon Irish potato in-
cubated for forty-three days at 37° C. showed a few scattered,
rather poorly staining, partially autolyzed acid fasts in pure
culture, whereas a comparable smear from the same amount of
inoculum in a slant of glycerin ovomucoid yolk agar incubated
for the same time at 37° C, showed somewhat larger numbers
of brightly staining, typical acid fasts in pure culture, which dis-
appeared upon longer incubation. Continuous incubation at 37°
46,4 Oliver et al.: Mycobacterium leprae 615
C. for five to seven months of all of our original aerobic cul-
tures has not alone failed to reveal any evidence of multiplica-
tion of the implanted M. leprae, but in the extremest instances
of persistence of the microorganisms in aerobic cultures no acid
fasts were found after two months' incubation. Aerobic trans-
plants to the same medium, as well as to all other media employed
in our studies, made at weekly intervals and at the end of four
and six weeks' continuous incubation, from all original aerobic
cultures that showed a persistence of M. leprae in pure culture,
have consistently failed to reveal any evidence of multiplication
of the transferred M. leprae. Moreover, in aerobic transplants,
the numbers of M. leprae have consistently diminished, and in se-
cond and third transplants the acid fasts have entirely disap-
peared.
In all of the twelve cases studied, partial oxygen tension
cultures revealed a longer persistence of M . leprae, in larger num-
bers, and with more typical morphological and staining charac-
ters, than did the corresponding aerobic cultures. Usually, cul-
tures made anaerobic with pyrogallic acid and potassium hy-
droxide yielded a longer persistence of M. leprae in larger
numbers, in larger clumps and more vividly acid fast, than did
the corresponding partial oxygen tension cultures; in partial
tension cultures, M . leprae appeared somewhat shorter than in
anaerobic cultures. Acid fasts, both granular and solid forms,
were quite regularly found in the largest numbers if smears were
made from a fragment of blood or tissue implanted in the water
of syneresis, or on the surface of the medium just above the
water of syneresis. However, in no instance in any of our
partial tension and anaerobic cultures, originals as well as in
transplants, was observed any evidence of microscopic, or un-
disputable microscopic, multiplication of M. leprae. As in the
aerobic series, continuous incubation was employed at 37° C.
and at room temperature for periods ranging from five to seven
months. In both the partial tension and in the anaerobic series,
the best and longest persistence of M. lepras was obtained on
glycerin ovomucoid yolk agar, glycerin bouillon saba wedges,
infant brain agar, and in one instance 4 per cent glycerin chick-
en bouillon sweet potato, at 37° C. In certain of the original
anaerobic and partial tension cultures on these media M. leprae
have been found in large numbers in pure culture after ninety-
six days of continuous incubation, the extreme instance being
one hundred fifty-eight days. First transplants have revealed
616 The Philippine Journal of Science 1931
still considerable, but smaller, numbers of M. leprae in pure
culture after twenty-four to ninety-six days incubation; and
second transplants have showed a persistence of still smaller
numbers of M. leprae in pure culture for shorter periods of time,
ranging from twenty-four to thirty-six days. Aerobic trans-
plants from partial oxygen tension and anaerobic original cul-
tures revealed a persistence of M. leprse in decreasing numbers
for a few weeks, with diminished acid-fastness, and their com-
plete disappearance within one month or less. On the other
hand, partial oxygen tension transplants from original partial
oxygen tension cultures, anaerobic transplants from anaerobic
original cultures, as well as partial oxygen tension transplants
from anaerobic cultures and anaerobic transplants from partial
oxygen tension cultures, all revealed a longer persistence, in
larger numbers, of more typical and more brightly acid-fast
bacilli.
In the case of our partial oxygen tension series of cultures,
originals as well as transplants, no demonstrable difference in
the persistence of M. leprse, nor in its numbers, was noted in
cultures incubated continuously with the same original tube of
B. subtilis or B. coli attached, and in those to which fresh tubes
of B. subtilis or B. coli were attached daily. Again, no signifi-
cant difference was observed in the persistence of M. leprse
when transplants were made from original cultures at weekly
intervals, at the end of four to six weeks' continuous incuba-
tion, and after two to four months of continuous incubation.
However, the longest persistence of M. leprx in large numbers
was observed by us in an original partial tension culture made
from case 9. This culture, made by planting a loopful of
bloody scrapings on the surface of a 4 per cent glycerin chicken
bouillon sweet potato wedge just above the glycerin, was in-
cubated continuously at 37° C. for eighty-eight days at partial
oxygen tension, with the original slant of B. coli attached. At
the end of this period, oxygen was admitted to the culture, ac-
cording to the method suggested by Wherry,4 by inserting a
needle through the connecting rubber tubing into the cotton
plug of the culture tube. The culture was then reincubated
at 37° C. for another seventy days, making a total incubation
period of one hundred fifty-eight days, at the end of which time
the culture was examined. A smear made from a fragment
of the dark brown inoculum revealed, on a five-minute search,
4Loc. cit.
46'4 Oliver et al.: Mycobacterium leprae 617
twenty clumps of vividly acid fast, granular, M. leprae in pure
culture. The majority of the clumps consisted of from four to
about thirty M. leprae, with a few scattered groups of two to
three members.
As illustrative of our anaerobic results, we might cite cer-
tain of our cultural findings on case 6. At the time of making
the original cultures from this case, direct smears revealed
M. leprae in relatively small numbers, an average of about ten
acid fast bacilli per oil-immersion field; the majority were in
small clumps of six to eight members. A platinum loopful of
bloody scrapings from the incised lesion was immediately planted
on the surface of a glycerin bouillon saba wedge, just above
the glycerin, and the culture was made anaerobic with pyro-
gallic acid and potassium hydroxide. At the end of twenty-six
days continuous incubation at 37° C, the tube was opened and
a smear was made from a small fragment of the implanted
bloody scraping. This smear revealed in pure culture, quite
large numbers of typical M. leprae, granular and solid forms,
averaging from thirty to forty per oil-immersion field; the ma-
jority of the acid fasts were in clumps of from fifteen to about
one hundred members. In as much as the direct smear from
the lesion had shown only about ten M. lepra? per oil-immersion
field, the majority in small clumps of six to eight members,
these cultural findings seemed somewhat encouraging. They
might be interpreted as suggesting that the implanted M . leprae
had undergone a certain amount of preliminary multiplication;
again, the results might be explained by assuming that the loop-
ful of inoculum used for this particular culture tube contained
a larger number of M . leprse than did the material used for
making the direct smear. Which of the two interpretations is
the correct one, we do not know; it seems to us, however, that
the second alternative is at least equally as valid as the first.
At any rate, the subsequent history of this original culture would
indicate that if a certain initial proliferation of M. leprse did
occur, this mulplication was not sustained. On examination of
the culture at the end of fifty days continuous anaerobic incuba-
tion at 37° C, a smear made from approximately the same-sized
fragment of implanted blood scraping revealed acid fasts in
smaller numbers, averaging from eight to ten per oil-immer-
sion field, and after ninety-six days of continuous anaerobic
incubation, no M. lepras were found. Analogous anaerobic and
partial oxygen tension cultures made on 6 per cent glycerin
ovomucoid yolk agar and on glycerin Irish potato wedges at
264209 6
618 The Philippine Journal of Science 1931
the same time as the original anaerobic saba wedge culture was
made, revealed only a few scattered acid fasts after twenty-six
days at 37° C, and at the end of fifty days' incubation no M.
leprae were found.
An anaerobic glycerin bouillon saba wedge transplant 1, was
made from the original anaerobic saba culture at the end of
twenty-six days' incubation at 37° C, an amount of inoculum
being employed that was approximately the same as in making
the smear. After forty-nine days at 37° C, this transplant 1
revealed in pure culture an average of about ten M. leprae per
oil-immersion field. The majority of the acid fasts were of
the granular type, but scattered, longer, thinner, solid forms
occurred ; clumps of from twenty to fifty plus brightly staining
acid fasts were found. Anaerobic transplant 2 on glycerin
bouillon saba, made from anaerobic saba transplant 1 at the
end of forty-nine days at 37° C, revealed on eleven days' in-
cubation at 37° C. only one granular acid fast; on twenty-five
day's incubation, as well as at the end of three months, no M.
leprae were found.
Anaerobic transplant 1 to a glycerin Irish potato wedge, made
from the original glycerin bouillon saba wedge after twenty-six
days of anaerobic incubation, revealed only a few, scattered,
feebly acid fast M. leprae in pure culture after forty-nine days'
incubation at 37° C, and by the end of ninety-five days' incuba-
tion no acid fasts could be found. Anaerobic transplant 1 to the
water of syneresis of a 6 per cent glycerin ovomucoid yolk agar
slant, made from the original anaerobic glycerin bouillon saba
culture after twenty-six days of incubation, revealed no acid
fasts at the end of forty-nine, sixty-three, and ninety days'
incubation. Partial oxygen tension transplant 1 on glycerin
Irish potato, glycerin bouillon saba wedges, and into the water
of syneresis of glycerin ovomucoid yolk agar slants, made from
the original anaerobic glycerin bouillon saba culture after it had
been incubated twenty-six days, revealed a few scattered M.
leprae in pure culture after forty-nine days at 37° C, but on
later examinations no acid fasts could be found.
SUMMARY
The longest persistence of M. leprae in pure culture, and in
large numbers, that was observed by us was one hundred fifty-
eight days. This occurred in an original partial oxygen tension
culture from case 9 on 4 per cent glycerin chicken bouillon sweet
46,4 Oliver et ah: Mycobacterium leprx 619
potato, incubated at 37° C. for eighty-eight days, with the orig-
inal tube of B. coli attached. At the end of this time, oxygen
was admitted according to the method suggested by Wherry,5
and the culture was reincubated for another seventy days.
In the remaining eleven cases, somewhat longer persistence
in pure culture of larger numbers of somewhat more typical
appearing M. leprx was obtained at 37° C. under anaerobic
conditions (pyrogallic acid and potassium hydroxide) than at
partial oxygen tension. Of the media employed by us, the most
favorable were glycerinized ovomucoid yolk agar slants
(Wherry medium), glycerin bouillon saba wedges, and infant
brain agar slants. In certain of such anaerobic cultures, M.
leprae have been found in pure culture, and in large numbers,
after ninety-six days of continuous incubation at 37° C.
The next best persistence of M. leprae in pure culture was
obtained in partial oxygen cultures on the same media at 37° C.
In certain of such cultures, M. leprx have been found in large
numbers after ninety-six days' continuous incubation, and in
one case after one hundred fifty-eight days. No demonstrable
difference was observed in cultures to which a fresh tube of
B. coli or B. subtilis was attached every twenty-four hours, and
in cultures to which the original tube of B. coli was allowed to
remain attached during the entire period of incubation.
The shortest persistence of M. leprae, in the smallest num-
bers, was obtained in our aerobic cultures, in which an irregular
persistence of M. leprx for from fourteen to forty-three days
was observed.
First, second, and third transplants, aerobic, partial oxygen
tension and anaerobic, have shown a progressive diminution in
the number of M. leprx, with a disappearance of acid fasts in
the third transplants. No demonstrable difference in the per-
sistence of M. leprx was observed when transplants were made
at weekly intervals, at intervals of three to six weeks, and at
intervals of two to three months.
The addition of cystine in 0.1 per cent concentration, of nor-
mal human serum, and of syphilitic serum appears to exert no
favoring action upon the persistence of M. leprx in cultures.
In none of our original cultures and transplants, which were
incubated at 37° C. for periods ranging from four to seven
months, did we obtain any macroscopic evidence of multiplica-
5 Loc. cit.
620 The Philippine Journal of Science 1931
tion of the implanted M. leprse, nor did we obtain any indis-
putable microscopic evidence of proliferation.
APPENDIX; DESCRIPTIONS OF CASES
Case 1 — Vicente Arriola.
Age 32. Male. Single. Filipino. Occupation, fireman. Family his-
tory for leprosy negative. First sign and symptom, anaesthesia on left
knee six months ago. Never received antileprotic treatment. Physical
condition good. Local lesions are numerous, small, pale pinkish, slightly
elevated macular patches all over trunk, buttocks, upper extremities, thighs,
the left and lower portion of the face. No nodules. Slight infiltrations
in face, ear lobes, and left knee.
Summary. — A case of moderate cutaneous, slight neural leprosy of six
months' duration. Cultures made from left malar eminence.
Direct smear. — M. leprae in small packets and singly; bacilli average
about 15 in number per oil-immersion field and are bright staining.
Case 2 — Claudio Quinto.
Age 40. Male. Married. Filipino. Occupation, quarryman. Family
history for leprosy negative. First sign and symptom, anaesthesia on both
legs, preceded by nodules, five years ago. Never received antileprotic treat-
ment. Physical condition fair. Local lesions are dry, wrinkled, macular
patches on trunk and thigh; extensive infiltrations on ears and face, less
extensive on trunk, buttocks, and extremities. Indurated, thickened skin
on legs and feet. Anaesthesia extensive on both extremities.
Summary. — A case of advanced cutaneous and advanced neural leprosy
of five years' duration. Cultures made from right ear.
Direct smear. — Enormous numbers of bright-staining M. leprae, the ma-
jority in large or small masses, but with many single acid-fast rods. The
number of bacilli averages over 100 per oil-immersion field.
Case 3 — Pablo Carpio.
Age 19. Male. Single. Filipino. No occupation. Family history for
leprosy negative. First sign and symptom, anaesthesia on forearm and
right elbow, six years ago. Never received antileprotic treatment. Phys-
ical condition fair. Local lesions are extensive; infiltrations on ear, face,
nipples, hands, and feet; no macules. Beginning nodular pigmentation
and taenia of the ear, medial aspect of palm, forehead, and feet; indurated,
thickened skin on legs.
Summary. — Advanced cutaneous and advanced nodular leprosy of six
years' duration. Cultures made from right ear and forearm.
Direct smear. — Enormous numbers of bright-staining M. leprae, the ma-
jority in large or small masses, but with many single acid-fast rods. Ba-
cilli average over 150 per oil-immersion field.
Case 4 — Juana Falcone.
Age 36. Female. Single. Filipino. No occupation. Family history
for leprosy negative. First sign and sympton was a slight numbness over
right leg one year ago. Never received antileprotic treatment. Physical
condition poor. Local lesions are macules, numerous on back, anterior
chest wall, and epigastrium. Infiltration diffuse on face, ears, fingers,
46,4 Oliver et al.: Mycobacterium leprae 621
and toes. Nodules, large and small, on elbows, forearms, buttocks, arms,
and lower extremities. Maculopapular areas on lower and upper extre-
mities.
Summary. — Advanced cutaneous and slight neural leprosy of one year's
duration. Cultures made from right ear lobe.
Direct smear. — Relatively small numbers of brightly staining M. leprae,
practically all single. Becilli average about 5 per oil-immersion field.
Case 5— €regorio Jabines.
Age 44. Male. Widower. Filipino. Occupation, merchant. Family
history for leprosy negative. First sign and symptom was one, large
reddish macule on abdominal wall two years ago. Never received anti-
leprotic treatment. Condition fair. Local lesions are macules on face,
ears, chest, abdomen, back, lower and upper extremities, and nape of neck.
Nodule on ear; no infiltrations.
Summary. — Moderate advanced macular leprosy of two years' duration.
Cultures made from arm.
Direct smear — M. leprse in moderately large numbers averaging about 30
per oil-immersion field. Majority of bacilli are single, with occasional
small masses of four to eight members. M. leprse stains more feebly than
in cases 1 and 4.
Case 6 — Rosa Musni.
Age 14. Female. Single. Filipino. No occupation. Family history
for leprosy negative. First sign and symptom was anaesthesia of forearm
one year ago. Never received antileprotic treatment. Condition good.
Local lesions are macules on cheeks, ears, chest, abdomen, and right fore-
arm. Infiltrations on ear. No nodules. Ichthyosis, both legs.
Summary. — Moderate cutaneous and slight neural leprosy of one year's
duration. Cultures made from left ear.
Direct smear. — M. leprse in relatively small numbers, averaging about
10 per oil-immersion field. Majority are in small masses of six to eight
members. M. leprse stains slightly more brightly than in case 5.
Case 7 — Laynes.
Age 18. Male. Single. Filipino. Occupation, laborer. Family his-
tory for leprosy negative. First sign and symptom was redness of face
and thickening of nose and ears three months ago. Never received anti-
leprotic treatment. Condition fair. Local lesions are macules, big and
depigmented, on trunk, arms, and thighs. Infiltration extensive on face
and ear. No nodules.
Summary. — Macular, moderately advanced cutaneous, and slight neu-
ral leprosy, of three months' duration. Cultures made from left ear.
Direct smear.— M. lepras in moderate numbers, averaging about 35 per
oil-immersion field. Majority are in masses of fifteen to twenty-five mem-
bers; bacilli stain brightly.
Case 8— Yep Hensr.
Age 25. Male. Single. Chinese. Occupation, carpenter. Family his-
tory for leprosy negative. First sign and symptom were reddish, anaes-
thetic patches on internal aspect of right knee about one and a half years
ago. Never received antileprotic treatment. Condition good. Local le-
622 The Philippine Journal of Science 1931
sions are macules on face, ears, chest, abdomen, back, nape of neck, and
upper extremities. No infiltration; no nodules.
Summary, — Moderate macular leprosy of about one-half year's duration.
Cultures made from cheek.
Direct smear, — M. leprae in relatively small numbers, averaging about
15 per oil-immersion field. Single acid fasts predominate, with occasional
clumps of eight to ten members. M. leprae stains rather feebly.
Case 9 — Leopoldo Duque.
Age 16. Male. Single. Filipino. Occupation, farmer. Family his-
tory for leprosy reveals a leprous father. First sign and symptom were
whitish areas on right knee about one year ago, after recovery from
measles; also large areas of same character on abdomen. Never received
antileprotic treatment. Condition fair. Local lesions are macules on
cheeks, abdomen, back, lumbar region, right arm, and anterior aspect of
thighs. Infiltrations in alae nasse and cheeks. No nodules. Ulnar and
common perineal nerves thickened.
Summary. — Moderate macular and moderate neural leprosy of about
one year's duration. Cultures made from ear.
Direct smear. — M, leprae in relatively small numbers, averaging about
twelve per oil-immersion field. Majority of bacilli are single, with oc-
casional clumps of eight to twenty members. M. leprae stains brightly.
Case 10— Feliciano Duque.
Age 42. Male. Married. Filipino. Occupation, farmer. Family his-
tory for leprosy reveals a leprous brother-in-law. First sign and symp-
tom were numbness over both feet about fourteen months ago. Never re-
ceived antileprotic treatment. Condition fair. Local lesions are macules
on cheeks, chest, abdomen, back, lumbar region, both arms, buttocks, and
anterior aspect of thighs. Infiltrations on both ears and cheeks. No no-
dules. Ulnar, great auricular, and common perineal nerves thickened.
Summary. — Moderate macular and moderate neural leprosy of about
fourteen months' duration. Cultures made from ear.
Direct smear. — M. leprae in small numbers, averaging about two per
oil-immersion field. Majority are single, with very occasional clumps of
four to eight members. Bacilli stain brightly.
Case 11— Jose Cabie.
Age 17. Male. Single. Filipino. Occupation, farmer. Family his-
tory for leprosy reveals a leprous mother and brother. First sign and
symptom were redness on cheek and thickening of ears ten years ago. Re-
ceived antileprotic treatment, and several injections, several years ago.
Physical condition good. Local lesions are extensive macules and nodules
over face, ears, arms, and entire body, with ulcerations on legs. Slight
enlargement of ulnar nerve. Atrophy of digits.
Summary. — Moderately advanced cutaneous and advanced neural leprosy
of ten years' duration. Cultures made from ear, with tissue and blood
scrapings.
Direct smear. — M. lepras in moderate numbers, averaging about twenty
per oil-immersion field. Majority are in loose or tight clumps of twenty-
five to fifty members. Acid fasts stain brightly.
46,4 Oliver et ol.; Mycobacterium leprae 623
Case 12— Valeriano Cabie.
Age 18. Male. Single. Filipino. Occupation, farmer. Family his-
tory for leprosy reveals a leprous mother and brother. First sign and
symptom were reddish areas on face and body twelve years ago. Received
antileprotic treatment for one month three months ago. Physical condi-
tion good. Local lesions are extensive; macules and nodules over face,
ears, arms, and entire body, with ulcers on legs.
Swnmary. — Advanced cutaneous and neural leprosy of twelve years*
duration. Cultures made from ear, with tissue and blood scrapings.
Direct smear. — M. leprm in very large numbers, averaging about one
hundred per oil-immersion field. Majority are single, but there are many
loose clumps of eight to sixteen members, and tight clumps of twenty to
fifty members. Acid fasts stain brightly.
ILLUSTRATION
Plate 1
FlGS. 1 and 2. Smear from a 158-day growth of Mycobacterium lepras.
625
OtlYEE : IT al. ; Mycobacterium L1PBJS,]
.V
[Philip, Jooen. Sci., 48, No, 4
fiff
;# ,?
1
*lL
* If
■ 7 m
r.0
PLATE 1.
SOMATIC SEGREGATION IN DOUBLE HIBISCUS
AND ITS INHERITANCE 1
By N. B. Mbndiola
Of the Department of Agronomy, College of Agriculture
University of the Philippines
THREE PLATES AND ONE TEXT FIGURE
INTRODUCTION
The study of somatic segregation is important genetically for
at least two reasons. One reason is that many horticultural
varieties of plants have their origin in that type of somatic
segregation which is called bud mutation. A second reason is
that a somatic segregation may or may not be sexually heritable ;
hence offering no little difficulty to a geneticist who is studying
the inheritance in crosses in which one of the parents or the two
used arises as bud sports. An example may help in explaining
this point. I have in my plant collection a variety of cassava
(Manihot utilissima Pohl) which is highly ornamental because
the area comprising the center and base of the lobes of its leaf
is yellow when the leaf is young and white when it is old. This
variety came from a bud sport of another cassava which like
most plants of Manihot utilissima has solid-green leaves. I
have grown about a hundred seedlings of this ornamental cassava
and every one of them has solid-green leaves. Knowing its
vegetative origin it is of course easy to understand why the
ornamental character is not heritable sexually. But let us sup-
pose that we did not know its origin and that this ornamental
variety had been crossed for purposes of Mendelian study with
a variety with solid-green leaves. The results of such a cross
would be highly confusing for we would be looking in the F2
offspring for a character which would not be there. A con-
fusion arising in this way might be avoided by a study of selfed
seedlings of the parents used in crossing, but this is only possible
1 Experiment Station contribution No. 750. Read before the Los Baiios
Biological Club February 26, 1931. Received for publication June 17,
627
628 The Philippine Journal of Science 1931
in case of varieties that are self-fertile and self-compatible.
If they happen to be self -sterile and self -incompatible then only
a knowledge of their vegetative parentage could aid a geneticist
in explaining the confusing results obtained in hybridization
work involving such somatic segregation.
OBJECT OF THE PRESENT WORK
The object of this paper is to report three types of somatic
segregation that have occurred in "double" varieties of Hibis-
cus rosa-sinensis Linnaeus and the results of crosses made in-
volving flowers from these somatic segregates.
REVIEW OF LITERATURE
As early as 1913 Wilcox and Holt (1913) reported somatic
segregation in flowers of double hibiscus. They reported that
"on the Double Salmon there are occasionally dark red double
flowers, and the Double Yellow now and then bears a regular
double flower half yellow and half salmon, or occasionally flowers
which are of salmon color throughout."
The first published report on somatic segregation in Hibiscus
in the Philippines was made by Mendiola and Capinpin in 1923.
They then reported a case of a bud sport consisting of a branch
of a pink variety of H. rosa-sinensis producing white flowers;
and another case of a branch of a red variety producing pink
flowers. They also reported cases of plants with entire leaves
producing lobed leaves usually at the base of the plant. This
case of dimorphism was subsequently studied by the senior
author and in a paper published in 1926 he reported that such
presence or absence of lobing in the leaves occurred at the ju-
venile stage of a plant arising either as a seedling or as a cutting
and that the presence of lobes in juvenile leaves of Hibiscus
rosa-sinensis is a simple dominant over absence of lobes.
While I was in Java in 1927, 1 received a letter from Dr. Jean
Schweizer, botanist of the Besoekishch Experiment Station in
that country, calling my attention to the occurrence in his gar-
den of a bud variation in Hibiscus consisting of a simple red
flower which had been produced by a plant of the Double Salmon
variety. Doctor Schweizer described the simple flower as in no
way different from the common hedge Hibiscus. By the com-
mon hedge Hibiscus I refer to the variety that is common in
the Philippines, which I have been calling Native Red Single.
46> 4 Mendiola: Double Hibiscus 629
TIME AND PLACE OF THE WORK
The work here reported which covered a period of more than
one year, was started December 30, 1929, and the experiments
were performed both in the author's plant-breeding garden and
in the plant-breeding nursery of the College of Agriculture, Los
Baiios, Laguna.
OBSERVATIONS AND EXPERIMENTS
CASES OF VEGETATIVE SEGREGATION OBSERVED
A periclinal chimera in a flower of the Double Salmon. — A
case was found in whIJi a Double Salmon plant produced a
flower which was similar to that of a Double Red except that a
number of the petals in the center of the flower remained salmon
in color. This would appear to be a case of a periclinal chimera
and suggests that the Double Red might have originated as a
bud sport of the Double Salmon. The flower showing the chi-
mera was not used in any pollination work as it was severed
from the plant when it was given to the author by Professor
Herbert, formerly of our Plant Physiology Department.
A bud sport consisting of Double Carmine produced by Double
Rose. — If one examines a flower of a Double Rose hibiscus he
finds that its "eye" is carmine in color. It apears that through
some somatic change during the early history of a bud, a branch
can arise which produces flowers all carmine in color instead
of being rose with a carmine eye. Such a branch has been pro-
duced, and by its propagation by cuttings it has made possible
the existence of the variety which we call Double Carmine.
It is quite possible that the first case is a manifestation of
the same phenomenon as the second except that in the former,
for some reason, the segregation was not completed.
Simple flowers from double varieties. — The production of a
simple flower by a double variety has been observed on one
branch of a plant of the Double Carmine (see Plate 1) and four
branches of a plant of the Double Rose. Two simple flowers
were produced by the Double Carmine plant ; one was produced
December 30, 1929, and the other January 5, 1930. One branch
of the Double Rose produced one single flower, in August, 1930.
Two other branches, January 29, 1931, produced simple flowers.
One of these branches produced one simple flower, and the other
branch produced one simple flower and one flower with two
whorls of five petals each. These three flowers were excep-
630
The Philippine Journal of Science
1931
Fig. 1. A periclinal chimera shown by a flower of the Double Salmon hibiscus.
tionally large, measuring 17 centimeters across with pistil about
9 centimeters long. The simple flowers produced by the double
varieties were used in pollination experiments which are de-
scribed later in this paper. For convenience, the simple flowers
from Double Carmine are henceforth called Mutant Simple Car-
mine and those from Double Rose, Mutant Simple Rose.
EXPERIMENTS
Self-pollination. — The two single flowers produced by the
Double Carmine were self-pollinated the days they were pro-
duced to determine if they were self -compatible and to find out
if and how simpleness occurring as a bud segregation in a double
variety as well as the carmine color arising from a rose variety
with carmine eye were heritable. The same thing was done
with the two simple flowers and with the flower with two rows
of petals produced by the Double Rose January 29, 1931.
Cross-pollination. — Besides self-pollinating the simple flowers
produced by the Double Carmine, they were crossed with nine
simple varieties using the former as the source of pollen. The
amount of pollen, coming as it did from only two flowers, was
quite limited and it was not possible to use as female more than
46> 4 Mendiola: Double Hibiscus 631
a total of eighteen flowers of the nine simple varieties men-
tioned.
The two single flowers produced by Double Rose January
29, 1931 were used as male and crossed with fifteen flowers of
seven single varieties.
RESULTS OF EXPERIMENTS
Of self '-pollination.— -The simple flowers produced by the Dou-
ble Carmine and the Double Rose failed to ripen pods upon self-
pollination, suggesting the probability of self-incompatibility
in this case of somatic segregation.
Of cross-pollination. — Of the nine different crosses made be-
tween the Mutant Simple Carmine and Single varieties, involving
eighteen flowers on the female side, only one succeeded ; namely,
that with the variety 19-Pink 12396-104. The results of cross-
pollination between the Mutant Simple Rose and other varieties
are not reported in this work.
The hybrids obtained. — From the successful cross eight plants
were obtained. They were given pedigree Nos. 5333, 5334,
etc., up to 5340 (see Plate 2). Table 1 gives a brief description
of the hybrids. Hybrid 5340 is shown in Plate 3, fig. 1, and
hybrid 5333, in Plate 3, fig. 2. It will be noted in Table 1 that
of the eight hybrids produced, four turned out to be double
and four, single, suggesting a 1 : 1 ratio and the heterozygosity
of one of the parents. While no two of these eight hybrids
are exactly alike and none of them is exactly like any other
existing variety, most of them are not enough different to war-
rant their being considered novelties. However, the case is dif-
ferent with No. 5339, which has light French vermilion corollas.
As there is no known vermilion double hibiscus, this hybrid,
which we will now call Double Vermilion, constitutes an entirely
new horticultural variety.
It may be of interest to note here certain habits of some of
the hybrids. Let us take No. 5333. This hybrid has a habit
which may be called telescopic production of two successive
flowers on one peduncle. About ten days after a flower is pro-
duced normally on one flower stem, instead of the evidently
empty pod falling off, another flower bud begins to come out of
it. The bud develops and opens slowly for about twenty days,
then it falls off — the flower never opens completely. The second
flower is completely sterile, and the normal is fertile in both
sexes. A plant with a similar habit was reported by Wilcox and
Holt (1913).
632
The Philippine Journal of Science
1931
Table 1. — Description of hybrids between simple flowers of Double Carmine
and a simple variety, 19-Pink 12396-104, and of the parents.
Double Car-
Simple flow-
ers from
19-Pink
Hybrid No.—
mine.
Double Car-
mine.
12396-104.
5333
5334
5335
Corolla
Multiple
Single
Single
Multiple
Single
Single.
Color o! corolla. _
Carmine
Carmine
Yellow-.
Light
Yellow- .
Light
French
vermi-
vermi-
lion.
lion.
Color of eye
Carmine
Carmine
No eye
Dark car-
No eye
Carmine.
mine.
Hybrid No. —
5336
5337
5338
5339
5340
Corolla
Single
Multiple
Carmine
Multiple
Carmine
Single
Light French
Multiple.
Rose.
Color of corolla..
Light orange
with ver-
vermilion.
milion
veins.
Color of eye
Carmine-
Dark carmine
Carmine.
Hybrid 5338 produces flowers which either never open com-
pletely or open only after about two weeks from the time the
petals begin to appear. As in the case of the telescopic flower
of No. 5333 the flowers of No. 5338 stay on the stem for a long
time. The persistence of the flower on the stem and the in-
ability to open completely appear to be correlated in the case of
these two hybrids. What significance this correlation may have
as a general biological phenomenon arouses our curiosity.
DISCUSSION OF RESULTS
Because of the accidental nature of the somatic segregation
reported and used in this work and the very limited number of
flowers which could be used in the pollination experiments and
the consequent small number of hybrids which could be obtained,
the results presented in this paper necessarily do not have the
reliability which more ample data possess. However, as the
accident might not happen again for many years or might never
happen again, it seems excusable to report these results at this
time. Besides, limited though their source is, they neverthe-
less suggest a number of interesting biological conclusions.
46' 4 Mendiola: Double Hibiscus 633
Origin of doubleness and of double varieties.— The term dou-
bleness when used to describe a flower refers to the number of
its corollas, and means either two corollas or more than two.
The corollas may be arranged as concentric whorls or each
may occur with its own center and outside the other. Consider-
ing the kinds of vegetative segregation reported in this paper,
it may be considered as certain that some of our double varieties
are color segregates of other double kinds, while soime of our
simple varieties arose as bud sports of others which bear mul-
tiple corollas. It is interesting to note here that in the case
of Hibiscus syriacus Linnaeus the Double and Simple white
varieties are similar in practically all respects, except that one
produces double flowers and the other simple flowers. The same
is true with the Double Lilac and Simple Lilac. Furthermore,
the white varieties are similar to the lilac, except in color. All
of these similarities suggest common vegetative parentage for
all these four varieties of H. syriacus, sexual parentage being
highly improbable as none of them is self -fertile.
When we attempt to explain the origin of doubleness in Hi-
biscus we find ourselves treading on more theoretical grounds.
The habit already cited of hybrid 5333, of bearing two successive
flowers on one flower stem seems to me to betray the secret of
the origin of doubleness. Hybrid 5333 is a very prolific flower
bearer. These two characteristics of No. 5333 suggest that
doubleness has its origin in heterosis, or hybrid vigor. It is
possible that natural crossing between two previously existing
simple varieties produced hybrid vigor in the hybrid. The
hybrid vigor resulted in the capacity of the hybrid to produce
a much greater number of flowers than either one of its parents.
Some of these flowers are produced simultaneously on one flower
stem, resulting in doubleness. Sometimes a flower is produced
much earlier, resulting in the production of a simple vegetative
segregate on a double variety or sometimes the flowers coming
out on one stem are produced successively, resulting in the teles-
copic appearance as exhibited by hybrid 5333.
Appearance of rose color of petals in a cross between carmine
and yellow. — It will be recalled that the hybrids reported in this
experiment are F1 progeny of a cross between a variety with
carmine petals and another with yellow petals. Among these
hybrids is one, No. 5340, that produces rose petals. The ap-
pearance of the rose color may best be explained by recalling
that the carmine parent arose as a bud sport of a rose plant.
264209 7
634 3T/i0 Philippine Journal of Science 1931
It seems probable that the carmine sport, while carmine pheno-
typically, retains a determiner for rose color which reappears
when conditions become favorable.
Because of the small number of hybrids raised, it is not ad-
visable to consider the results of the cross statistically. It may
be pointed out, however, that according to the assumption made
above, there should result a greater number of carmine individ-
uals than either yellow or rose, and this is exactly what hap-
pened, as shown by Table 1.
Before concluding the discussion of the appearance of rose
in a cross between carmine and yellow, it is well to point out
again that a knowledge of the vegetative origin of a variety
is bound to prevent us from adopting a wrong explanation of the
unexpected appearance of a character among hybrids. Taking
as an example the appearance of rose in a cross between carmine
and yellow already described, it is quite likely that had we not
known that the carmine parent arose from a rose variety, we
would be explaining the appearance of the rose color on the
basis of complementary factors, an explanation that is not the
best at all to offer at the present time.
Simple on double not involving mutation of a factor. — It has
been pointed out that the hybrids reported in this paper con-
stitute the first filial generation of a cross between a simple
flower produced on a branch of a double variety and another
simple flower of a simple variety. The fact that nearly half
of the hybrids turned out to be double shows that the appearance
of the simple flower on a branch of a double variety was not
due to a mutation of a factor for doubleness into that for sin-
gleness, this conclusion being supported by the additional evi-
dence that the branch that once produced the simple flower
continues to produce double flowers.
Inheritance of doubleness. — Table 1 shows that of the eight
hybrids obtained four were single and four were double, sug-
gesting that one of the parents was homozygous and the other
heterozygous. In the past I have performed numerous crosses
between simples and as no doubles have been produced this way
it may be concluded that simpleness is a homozygous recessive
while doubleness is a heterozygous dominant. Dd may represent
doubleness and dd simpleness. Dd X dd will result in 2Dd : 2ddf
or 50 per cent double and 50 per cent single. The somatic seg-
regation of a single flower from a double may be explained by
D of Dd changing to d.
46' 4 Mendiola: Double Hibiscus 635
Inheritance of the somatic segregate carmine.— In all the
crosses which we have made in the past between rose and yellow,
no carmine ever appeared among the hybrids. Furthermore,
sellings made of either rose or yellow never produced carmine.
These facts indicate that in the carmine hybrids produced in the
cross Mutant Simple Carmine X 19 Pink 12396-104, which
cross may for convenience be stated as carmine x yellow, the
carmine was due to the mutant carmine and not to any recessive
carmine combined with the pink of the original rose parent of
the carmine segregate. These facts also prove that while car-
mine appeared as a somatic segregation, the change affected the
germplasm with the result that it became heritable sexually.
SUMMARY
1. This paper reports three cases of somatic segregation found
in double varieties of Hibiscus rosa-sinensis Linnaeus in the
Philippines; namely, (a) a periclinal chimera in a flower of
the Double Salmon, (6) a branch of a Double Rose producing
double carmine flowers and serving as the origin of our Double
Carmine variety, (c) single flowers being produced by branches
of Double Rose and Double Carmine varieties.
2. Through the production of the simple flowers by the Double
Carmine variety, it was possible to cross this with a simple
yellow variety. Among the hybrids produced by this (double
carmine X simple yellow) cross there was a carmine plant and
a rose individual, showing that while the carmine sport is car-
mine phenotypically, it retains the rose determiner.
3. The carmine somatic segregate was found to be sexually
heritable.
4. The somatic segregation consisting in the production of
simple flowers by a double branch does not seem to have in-
volved the mutation of a factor.
5. Attention has been called to the fact that a somatic se-
gregation which does not affect the germplasm and is not, there-
fore, sexually heritable is likely to confuse the results of a given
cross in which one of the parents is a variety that has originated
as one such vegetative segregate. It is quite likely that the
failure of a character of a given parent to appear in any of
its hybrid offspring may be traced to this cause. This makes
it important that we know the vegetative origin of our horticul-
tural varieties or clons.
636 The Philippine Journal of Science
LITERATURE CITED
Mendiola, N. B., and J. M. Capinpin. Breeding ornamental hibiscus.
Philip. Agr. 11 (1923) 217-230.
Mendiola, N. B. A Manual of Plant Breeding for the Tropics. University
of the Philippines, Manila (1926) XXIII + 360 with text figures and
plates.
Wilcox, E. F., and V. S. Holt. Ornamental hibiscus in Hawaii. Hawaii
Agr. Exp. Sta. Bull. 29 (1913).
ILLUSTRATIONS
Plate 1
A Double Rose hibiscus plant showing in the left circle a normal double
flower and in the right circle the simple flower which arose as a
vegetative segregate.
Plate 2
Hibiscus (19 Pink-12396-104 X Mutant Simple Carmine) Fi.
Plate 3
Fig. 1. Hibiscus hybrid 5340, showing its double flower.
2. Hibiscus hybrid 5333, showing its double flower.
TEXT FIGURE
Fig. 1. A periclinal chimera shown by a flower of the Double Salmon hi-
biscus.
637
Mbndiola: Douili Hibiscus,]
(.Philip. Journ. Scl, 46, Mo, 4,
PLATE 1,
i
h
<
""XsL
Mbndtola: Double Hibiscus,]
[Philip. Jouhn. So., 46, No. 4.
PLATE 3,
DAYTIME RESTING PLACES OF ANOPHELES
MOSQUITOES IN THE PHILIPPINES
FIRST REPORT l
By Paul F. Russell
Chief, Malaria Investigations, Bureau of Science, Manila
FOUR PLATES
INTRODUCTION
Referring to the resting places of anophelines, Boyd(la) in
his very useful textbook on malariology summarizes as follows
the places where one should look for the adults of Anopheles in
the daytime:
1. Inside houses, especially the sleeping rooms, the darker corners,
the ceiling, the wall behind furniture and pictures, dark clothing, or under-
neath furniture.
2. Underneath buildings. On leeward side of chimney bases and on
windward side of joists.
3. In stables, pigsties, chicken houses, kennels, hutches, etc. when oc-
cupied.
4. In porches and verandas.
5. In privies.
6. Under bridges and in culverts.
7. On the shady side of road-cuts, stream and ditch margins having pre-
cipitous banks; in open wells, under ledges, and in caves.
8. On weed grown piles of brickbats and stones.
9. In the interior of hollow trees, or in the spaces between buttress-
ing roots.
10. On the underside of leaves in dense thickets, or in clumps of shrubs
and annual plants, perhaps in clumps of high grasses as well.
11. In cracks in the ground.
Hehir(2) referring to the adult anophelines of India, writes
as follows:
Out-houses, bath-rooms, damp go-downs, shaded verandahs, cow-sheds,
coach houses, garages, stables, unoccupied thatched houses, with dirty
aThe International Health Division of the Rockefeller Foundation, of
which the author is a field director, is cooperating with the Bureau of
Science of the Philippines in malaria investigations. Mr. Domingo San-
tiago, field inspector of malaria investigations, made the routine catches
reported in the tables.
639
640 The Philippine Journal of Science 1931
soot-covered walls, are special day resorts for anophelines. In a native
village or bazaar select huts that are near a pond, water channel or other
breeding place of mosquitoes. Some favour the thatch beneath the eaves
of huts and houses and require a ladder to reach them. Anophelines are
rarely seen on whitewashed walls in the daytime and seldom at night.
They may often be found in holes in walls, or in the corners of rooms,
under beds and tables, in cupboards, on dark clothes in rooms; behind
pictures, doors, furniture, in open fire-places and chimneys, in lavatories;
under porches and in sheds, under bridges and culverts, in wells and un-
screened cisterns; they are fond of hiding in old boots (Wellingtons and
polo boots especially) , on saddles — leather seems to attract them. We may
see them at night wandering about the shady side of the mosquito net.
The wooden rafters of thatched houses are a favourite retreat. In such
shady places they are readily captured as they are probably asleep, and it
is usually easy to place the butterfly net or mouth of the test-tube quickly
and quietly over them. The distribution of some species of Anopheles is
very local, hence it is necessary to examine as many likely places as
possible.
From these descriptions one might imagine that it is never a
difficult matter to find adult anophelines in the daytime in
localities where abundant breeding is known to exist. But as
a matter of fact malaria surveys in the Tropics have frequently
in the past been handicapped because adult Anopheles mosqui-
toes, especially of the species carrying malaria, could not be
captured in their daytime resting places. These shelters have
often defied careful search by skilled field inspectors.
Boyd(l&) refers to this fact and comments that Oriental ano-
phelines appear to be much less inclined to linger about dwellings
than do their relatives elsewhere.
MacGregor(3) in his excellent manual for mosquito surveys
cautions that it is well to note that certain species of anophelines
which enter houses to bite the inhabitants rarely remain in the
house after they have fed. They always attempt to get out of
doors immediately after feeding. Consequently, the presence
of these species is not to be detected by a daytime search.
Hackett,(4) who has had wide experience with anophelines in
various parts of the world, wrote after a visit to the Orient, as
follows :
I had no idea until I visited the Far East how difficult it is to lay
hands on the adults of most of the principal malaria carriers.
LITERATURE
Without attempting an exhaustive survey of the literature
several references may be cited as of interest in connection with
this subject. These make it obvious that uniformity of results
46» 4 Russell: Resting Places of Anopheles 641
has not been the rule. In India, for example, it has not always
been a difficult matter to make daytime catches of adult ano-
pheline mosquitoes. Refer again to Hehir.(2) Also note a
report by Christophers (5) of some malaria surveys in 1925.
In this report Christophers writes as follows :
Anopheles in the houses at the time of my visit, though no longer very
numerous, were to be obtained without great difficulty . . . Adult ano-
pheles at Manharpur were fairly abundant in cow-sheds, etc., at the Ba-
bus' old quarters.
Other Indian references could be cited which indicate that
adult Anopheles have been caught in daytime resting places in
the course of malaria surveys; but there have been real diffi-
culties, for Christophers, Sinton, and Co veil, (6) in their guide
for malaria surveys, comment as follows :
If catches of adults made in the houses are carefully and critically
examined in relation to the breeding places, and other catches of adults
made in the open, a very great deal may be learnt about the behavior of
Anophelines in particular circumstances. This work however has seldom
been attempted and there is consequently a large field for enquiry on
such lines.
In Ceylon only recently have daytime searches for adult Ano-
pheles been successful. For example, James and GunasekaraC?)
reported that adult anophelines were scarce and difficult to find.
Barnes and Russell (8) wrote, "It has generally proved difficult
to find the resting places of anopheles mosquitoes within village
houses." Carter (9) in his very complete report on malaria and
anopheline mosquitoes in Ceylon wrote:
Adult Anopheles were not always abundant even although the time
chosen for the visit appeared suitable, and on several occasions consider-
able difficulty was experienced in obtaining what were relatively small
numbers.
It was therefore notable when Carter and Jacocks(lO, ll) by
paying particular attention to the matter were able to find be-
tween 9 a. m. and 4 p. m. considerable numbers of Anopheles
resting on walls and hangings of village huts, small bungalows,
and coolie barracks in various localities in Ceylon. As a result
they were able to prove the importance of A. culici facies as a
malaria carrier in Ceylon by actual sporozoite findings in wild-
caught specimens, something which had never been done con-
clusively before. Plate 1, fig. 1, shows the type of coolie
barracks in which A. culicifacies were caught in the daytime, as
demonstrated to me in November, 1929.
642 The Philippine Journal of Science 1931
In British Malaya, while there have been reports of daytime
catches of Anopheles mosquitoes, not many records are to be
found. In the 1919 report of the Malaria Bureau (12a) there
is reference to 15 adults caught in thirteen days search at Per-
hentian Tinggi. In the 1922 report (126) it is noted that 10,327
adults had been caught in houses but apparently none were A.
maculatus, the chief malaria-carrying species. In the 1923 re-
port (12c) there is a record of 342 adult mosquitoes caught on a
Johore estate in the houses. Of these 335 were A. maculatus
and 11.5 per cent of the 199 dissected were found to be infected.
From personal experience in the Straits Settlements and in
Kedah, I know that it is difficult to find adult A. maculatus in
daytime resting places. In typical Malay houses they are sel-
dom to be found, and they do not rest under such houses. In
the darker and damper coolie barracks, or lines, they are more
apt to be found, but even here the catch of anophelines seldom
includes A. maculatus.
From the Dutch East Indies have also come reports both
of good catches of adult mosquitoes and of meager results.
Van Breemen(i3) reported catching large numbers of A. ludlowi
inside houses. In one locality adults were regularly caught,
although larvse could not be found. Swellengrebel and others(l4)
had no difficulty in catching A. ludlowi adults but speak of
''many other species which leave the house shortly after feeding
or which do not feed within the house. These species should
be caught in the daytime on plants or trees or under the house
or in the evening on man, cows or principally on buffaloes . . .
catching on buffaloes is a precious method to collect species not
to (be) found in houses." Schuffner and others(lS) had the
same experience and point out that A. ludlowi "is not found in
empty houses." These authors (15) speak of other anophelines
which after feeding "fly away again and hide themselves in
trees, shrubs, ditches or other cavities."
But even A. ludlowi has not infrequently been elusive. Brug
and Walch(l6) in Solo reported, for example, that "five coolies
under the supervision of a sanitary inspector could not catch
more than 6 anopheles in a kampong such as Tjinderedjo, where
at that time the parasite index of the children was 90, that for
the adults 65." The authors (16) themselves had no success al-
though they examined "dark corners and holes" and "crept
under bedsteads." They finally succeeded in catching Anopheles
46> 4 Russell: Resting Places of Anopheles 643
adults at night on buffaloes, although they had to offer a premium
for the catches.
Again in Tegal, Walch and Soesilod?) had to offer premiums
for adult anopheline catches which, although in one case averag-
ing forty-six, usually averaged less than two per catch.
Schuurman and Bokkel Huinink(i8) on the south coast of
Java caught fair numbers of anopheles adults "in and in the
neighbourhood of the houses.,,
Essed(l9) in reporting from Banjoewangi, Java east coast,
comments that "if these hiding places such as mosquito-nets
and dark recesses behind beds are not present, then one can
seek in vain in the native and other houses for Anopheles."
As to the western tropics the situation is the same as in the
East as regards catching adult anophelines. Some have reported
good catches. For example, Le Prince and Orenstein(20) wrote
of their experience in Panama as follows :
While no suitable hiding places except vacant houses were without
mosquitoes in the daytime, yet beyond the settled area none were
found. . . . Large numbers were collected under houses where the breeze
was sufficiently strong to make the lighting of a match difficult. These
inhabited houses were on posts from two to ten feet above the ground.
The dry weather ground-cracks under the houses were several inches deep
and the mosquitoes collected in them.
On the other hand Boyd and Aris(2i) wrote of their Jamaican
experiences as follows:
Searches made within houses during the day rarely yielded imagines,
even though large numbers could be caught at night in the vicinity from
a horse or mule as bait.
Stephens (22) in reporting a survey on a Venezuelan oil field
wrote as follows:
The search for anophelines in the native huts in the daytime was com-
pletely fruitless, and culicines also were very scanty. This condition was
in marked contrast to those observed by me in the neighborhood of Lake
Valencia, which I visited on my way home, where in the daytime, in the
verandah of a hut it was easy to collect numerous anophelines, embracing
three different species.
In Porto Rico Earle(23) relies not on routine daytime catches
but on traps baited at night with such animals as calves or
horses. He has been very successful with these traps. This is
true also of Manalang(23, 24) in the Philippines.
g44 The Philippine Journal of Science 1931
THE PHILIPPINES
In the Philippines as elsewhere in the Orient it is not an
easy matter to make routine daytime catches of anopheline adult
mosquitoes. Refer again to Hackett(4) who wrote:
In the seven days I spent in the [Philippine] Islands I did not catch
a single one [A. miniTrtus'] although a sporting colleague offered as high
as a peso [50 cents gold] apiece for adult specimens. At the same time
the larvae were abundant.
Manalang, who has made extensive and notable studies on A.
minimus (A. funestus) in the Philippines, writes: (25) "The adult
mosquito is typically 'wild' in that it is very seldom found in
the ordinary nipa house at night, much less in the day time."
The reason may sometimes be as suggested by Walker and Bar-
ber (26) that it is the custom of the people in the rural districts
to wash clothes and bathe in the streams, often in the early
morning or evening, thus affording ample opportunity to the
forest-loving anophelines. These observers, (26) however, made
fair daytime catches of adult mosquitoes in houses in Mindoro
and at Iwahig. In the latter place, a penal colony, the catches
were chiefly inside mosquito nets which had been badly adjusted.
They found only a few imagines along the banks of streams or
in crab holes. They suggested in their report (26) that certain
meteorological conditions influence the dispersal of adults.
Their negative results were in localities having at the time hot
and dry weather.
There have been no records of routine daytime catches of adult
Anopheles in the Philippines. Where these adults, particularly
A. minimus adults, go in the daytime has been and still is a ques-
tion. This paper gives some information, but much more in-
vestigation along this line is required.
The typical Filipino nipa house in rural areas is built high
off the ground. It is light, airy, and dry. It contains as a rule
very little furniture and no beds. Roofs may be either of tin
or thatching (Plate 1, fig. 2). At first glance it would seem
as though Anopheles mosquitoes would find ideal sheltering
places under such houses. In the Southern United States it has
been my experience, common to that of many others, that where
breeding is abundant it is more usual than not to find adults
of A. quadrimaculatus, the malaria carrier, under houses resting
on the sides of beams. In repeated searches in the Tropics I
have so far always failed to find mosquitoes in such places.
46, 4
Russell: Resting Places of Anopheles
645
Furthermore, Anopheles mosquitoes are rarely to be found in-
side such houses in the Philippines.
ROUTINE COLLECTIONS
During the last quarter of 1930 routine weekly catches were
attempted not only in certain houses (as shown in Table 1)
but also on the sides of a well (Table 2), along a stream bank
(Table 3), and in the cracks of a stone wall (Table 4). These
catching stations were all in or near Calauan, Laguna Province,
Luzon Island.
Table 1. — Adult Anopheles mosquitoes caught inside houses during routine
weekly collections October to December, 1930, Calauan. Only nighttime
catches are shown as no Anopheles mosquitoes were caught inside
houses by day. No traps used.
Species.
Anopheles subpictus (fresh water) _ .
Anopheles vagus (Philippine forms)
Total
Sex.
Total.
Male.
Female.
2
3
3
5
5
8
5
8
13
Table 2. — Adult Anopheles mosquitoes caught on the sides of an open well,
Masiit, October to December, 1980, in routine weekly collections,
[Catching time about ten minute3 once a week.]
Species.
Anopheles kochi (Donitz 1901)
Anopheles tessellatus (Theobald 1901)
Total
Sex.
Total.
Male.
4
36
Female.
8
56
12
92
40
64
104
The well used as a catching station is of the surface type hav-
ing no protection around the top except long grasses and low
bushes. The water level is about 7 feet below the ground sur-
face and the well is about 15 feet deep. The sides of the well
are of earth, and there are places where the top overhangs min-
iature caves in which adult mosquitoes are apt to be found
among exposed roots. Here it is darker, damper, and more
protected. The water of the well is not much used, and A.
tessellatus is breeding in it (see Plate 3, fig. 3, and Table 2).
646
The Philippine Journal of Science
1931
Table 3. — Adult Anopheles mosquitoes caught along a stream bank, Masiit,
October to December, 1930, in routine weekly collections.
[Catching time about thirty minutes once a week.]
Species.
Anopheles barter of ti var. pseudobarbirostris (Ludlow 1902)
Anopheles barbirostris (van der Wulp 1884)
Anopheles fuliginosus (Giles 1900)
Anopheles kochi (Donitz 1901)
Anopheles minimus (Theobald 1901)
Anopheles tessellatus (Theobald 1901)
Anopheles vagus (Philippine form)
Total
Sex.
Total.
Male.
Female.
3
3
6
6
7
13
1
3
4
4
8
12
126
272
398
36
56
92
52
93
145
228
442
670
Table 4. — Adult Anopheles mosquitoes caught along an old stone wall,
Calauan, October to December, 1930 in routine weekly collections.
[Catching time about fifteen minutes once a week.]
Species.
Anopheles barbirostris (van der Wulp 1884)
Anopheles hyrcanus var. sinensis (Wiedeman 1828)
Anopheles kochi (Donitz 1901)
Anopheles minimus (Theobald 1901)
Anopheles philippinensis (Ludlow 1902)
Anopheles tessellatus (Theobald 1901)
Anopheles vagus (Philippine form)
Total
Sex.
Male. Female.
2
0
3
6
1
2
75
1
1
9
9
0
26
64
110
Total.
3
1
12
15
1
28
139
199
In Plate 2 is shown the stream along which catches of adult
Anopheles were made. It is a typical A. minimus breeding
place of the Philippines. Plate 3, fig. 2, shows a catching sta-
tion. Here the stream bank is eroded and is overhung by veg-
etation, so that a darkened, damp, and sheltered resting place
is formed (see Table 3).
Another resting place for adult anophelines in the Philippines
is shown in Plate 4, figs. 1 and 2. This stone wall is in an old
cemetery about 1 kilometer from the nearest mosquito breeding
places. The wall is well shaded by high bushes, vines, and
trees. There are numerous large cracks and crevices in which
mosquitoes find darkened, sheltered resting places which are,
however, not very damp, in fact they seemed distinctly dry.
46, 4 Russell: Resting Places of Anopheles 647
From these findings it appears that routine daytime catches of
Anopheles imagines in the Philippines will have to include not
only human habitations or animal houses, but primarily natural
shelters such as cracks, crevices, and caves near or in the ground,
not necessarily near breeding places. Further searching in dif-
ferent types of houses and with nets among grasses and bushes
may reveal other sheltering places, but at the present time, for
the rural Philippines, the records in the tables of this report
may be taken as indicating the preferences of adult Anopheles
for their daytime resting places. For the catching of Anopheles
imagines in the Tropics trapping would seem to be the most
effective method, although traps still leave much to be desired.
SUMMARY
A brief review of the problem of catching adult Anopheles
mosquitoes in their daytime resting places in the Tropics is
presented. Some observations are given as to the situation in
the Philippines.
REFERENCES
1. Boyd, M. F. An Introduction to Malariology. Harvard University
Press, 1930. (a) Page 313. (b) Page 314.
2. Hehir, Sir Patrick. Malaria In India. Oxford University Press
(1927) 62.
3. MacGregor, M. E. Mosquito Surveys. Wellcome Bureau of Scientific
Research, London (1927) 235.
4. Hackett, L. W. Differences in the habits of anophelines which trans-
mit malaria in America, in Europe, and in the Far East. Southern
Med. Journ. 22, No. 4 (April, 1929).
5. Christophers, S. R. Two malarial surveys connected with industrial
projects in certain very highly malarious localities in India. Indian
Journ. Med. Res. 23, No. 2 (October, 1925) 355, 389-391.
6. Christophers, S. R., J. A. Sinton, and G. Covell. How to do a
malaria survey. Health Bull. 14. Malaria Bureau No. 6. Govt, of
India, Central Publication Branch (1928) 53.
7. James, S. P., and S. T. Gunasekara. Report on malaria at the Port
of Talaimannar. Sessional paper 34 (1913). Govt. Printer, Co-
lombo, Ceylon.
8. Barnes, M. E., and P. F. Russell. A programme for the control of
malaria in Ceylon. Sessional paper 9 (1926). Govt. Printer, Co-
lombo, Ceylon.
9. Carter, H. F. Report on malaria and anopheline mosquitoes in Cey-
lon. Sessional Paper 7 (1927). Govt. Printer, Colombo, Ceylon.
10. Carter, H. F., and W. P. Jacocks. Observations on the transmission
of malaria by anopheline mosquitoes in Ceylon. Ceylon Journ. Sci.
Sec. D. Med. Sci. 2 (1929) 67-86.
648 The Philippine Journal of Science
11. Carter, H. F. Further observations on the transmission of malaria
by anopheline mosquitoes in Ceylon. Ceylon Journ. Sci. Sec. D.
Med. Sci. 2, No. 4 (1930).
12. Federated Malay States, Malaria Bureau Reports, Kuala Lumpur.
(a) Vol. 1, Report VI (1919). (6) 1922 report, page 1. (c)1923
report.
13. Van Breeman, M. L. Malaria in Weltevreden and Batavia. Med. van
der Burg. Geneesk. d. in Ned. Indie, No. 2 (1919).
14. SWELLENGREBEL, N. H., W. SCHUFFNER, and J. M. H. SWELLENGREBEL DE
Graaf. The susceptibility of anophelines to malaria-infections in
Netherlands India. Med. van der. Burg. Geneesk. d. in Ned. Indie,
No. 3 (1919).
15. SCHUFFNER, W., N. H. SWELLENGREBEL, J. M. H. SWELLENGREBEL DE
Graaf, and Achmad Mochtar. On the biology of M. ludlowi in
Sumatra. Med. var den Burg. Geneesk. d. in. Ned. Indie, No. 3
(1919).
16. Brug, S. L., and E. W. Walch. Report of an investigation of a ma-
laria epidemic in Solo 1926. Med. van der dienst der Volksgez. in
Ned.-Indie (1927).
17. Walch, E. W., and R. Soesilo. Investigation of a malarial epidemic
in Tegal, during the first month of 1926. Med. v. d. dienst d. Volksgz.
in Ned. Indie, Part I (1927).
18. Schuurman, C. J., and A. Schuurman-ten Bokkel Huinink. A ma-
laria problem on Java's south-coast. Med. v. d. dienst d. Volksgez.
in Ned. Ind. (1929).
19. Essed, W. F. R. Malaria at Banjoewanji and the prospects of an
efficient species sanitation. Med. v. d. dienst d. Volksgez. in Ned.
Indie (1929).
20. Le Prince, J. A., and A. J. Orenstein. Mosquito Control in Panama.
G. P. Putnam's Sons, N. Y. (1916) 86.
21. Boyd, M. F., and F. W. Aris. A malaria survey of the Island of
Jamaica, B. W. I. Am. Journ. Trop. Med. 9, No. 5 (1929) 381.
22. Stephens, J. W. W. Malaria on a Venezuelan oilfield. Ann. Trop.
Med. & Parasit. 15, No. 4 (1921) 439.
23. Personal observation supplemented by personal communication.
24. Manalang, C. Does the amount of malaria depend on the number
of transmitting mosquitoes? Journ. Trop. Med. & Hyg. 34, No, 2
(1931).
25. Manalang, C. Notes on malaria transmission. Philip. Journ. Sci.
37 (1928).
26. Walker, E. L., and M. A. Barber. Malaria in the Philippine Islands.
Philip. Journ. Sci. § B 11 (1914).
ILLUSTRATIONS
[Photographs by the author.]
Plate 1
Fig. 1. Coolie barracks in Ceylon in which adult Anopheles culicifacies
were regularly caught in the daytime.
2. Typical rural Filipino houses.
Plate 2
A typical Anopheles minimus breeding place in the Philippines.
Plate 3
Fig. 1. Looking obliquely at the side of a well overhung by roots and
grasses. (See Table 2 and the text.)
2. A catching station for adult Anopheles minimus mosquitoes.
Plate 4
Fig. 1. Stone wall in old Calauan cemetery. (Time exposure.)
2. Close view of resting places of Anopheles on a stone wall. (Time
exposure.)
264209 8 649
Rksskll: Ukstinv. Planks of Anoimiki.ks.1
I Philip. Jofen. Sn., 46, No. 4,
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PLATE 1.
KtiKKKi.i.: Kkhtinc Places of Anoi-iiklks. I
ti'iULit1. Joi'KN. St'l., 4ti. No. 4.
PLATE 2.
HASKELL: ftesTlNfi Pl.ACKS OF A NO1MIKL10S. |
I Philip. Joifrn. Sn„ Ml, No. t.
PLATE 3.
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PLATE 4.
AVIAN MALARIA STUDIES, III
THE EXPERIMENTAL EPIDEMIOLOGY OP AVIAN MALARIA; INTRODUC-
TORY PAPER 1
By Paul F. Russell
Of the International Health Division, Rockefeller Foundation
TWO PLATES AND THREE TEXT FIGURES
INTRODUCTION
Although epidemiology as an ancient science dates back to
Hippocrates (l) it has had remarkably little experimented study.
Prior to Hippocrates epidemics were viewed entirely from a
supernatural or metaphysical standpoint. Witness, for example,
the Chaldaic malus annus, or evil year. In the Hippocratic
writings can be seen the first attempts to identify the natural
factors causing epidemics. Hippocrates gave little credence to
mysterious agents of the superstitious, but a conception of in-
fection was totally lacking, and so explanation of external causes
of epidemics, as based on Hippocrates'S observation, involved
careful consideration of the constitution of the atmosphere, of
cosmic influences, and seasons. This modest theory and Galen's
suggestive writings as to a distinction between exciting and
predisposing causes were unheeded until Guillaume de Baillou
(1538-1616), (2) physician to the Dauphin of Henri IV, reintro-
duced the idea that certain seasons and certain years are by
their peculiar innate nature subject to certain diseases. Bail-
lou has been called "the first epidemiologist of modern times/'
1 The first experiment reported in this paper was done in the Depart-
ment of Tropical Medicine, Harvard University Medical School, through
the courtesy of Prof. R. P. Strong and with the assistance of the Inter-
national Health Division of the Rockefeller Foundation. The second and
third experiments were done at the Bureau of Science, Manila, where the
author is chief of malaria investigations in which the bureau and the
division are cooperating. Misses Amparo Capistrano and Filomena Villa-
corta, of the staff of malaria investigations, assisted in the blood exam-
inations in the last two experiments. Mrs. Isabel Ramos, also of the staff,
assisted in handling the mosquitoes.
651
652 The Philippine Journal of Science 1931
Thomas Sydenham (1624-89) expanded this theory of genius
epidemicus, or "epidemic constitutions." Delving into accounts
of outbreaks of disease in London over a 25-year period, he
attributed great importance to the influence of season and
climate, not only in the origin of epidemics but even in deter-
mining predominant clinical manifestations of a given disease.
Smallpox under one epidemic constitution might be a very dif-
ferent matter as a disease and as an epidemic from smallpox
under another constitution. (3, 4, 5, 6)
Since Sydenham's time, except for an incorporation into the
concept of epidemic constitutions of the fact of infection by
demonstrable pathogenic organisms, there has been surprisingly
little progress in the development of the theory of epidemic
disease. Even to-day epidemics of malaria, influenza, plague,
cholera, and smallpox, for example, are in some phases as mys-
terious as they were to Hippocrates.
Present belief is in general as stated by Topley : "The origin
and spread of any epidemic of microbial infection depends upon
variations in the normally existing relations between living or-
ganisms; and the actual outbreak of disease, the occurrence of
clinically recognisable cases, is only the end-result of a progres-
sive disturbance of this normal equilibrium." (63) This being
true, Topley goes on to point out, the problems of epidemic
disease thereupon assume an aspect more biological than medical.
This last observation is especially true of malaria.
Not to mention the classical and historical methods, there are,
as Brownlee(i4) states, three ways in which the biological basis
of epidemics may be sought. There may be observation of the
mode of progress of the epidemic, in the first place, as it occurs
in nature or, in the second place, as during an experiment. In
the third place there may be an examination of accumulated
statistical information at our disposal.
Hippocrates, Baillou, and Sydenham followed the first way.
In their footsteps have come Maximillian Stoll (1742-1787), of
the old Vienna school; Lancisi (1655-1720), the Italian; and
more recently Hamer,(7-10) Crookshank,(H> 12) and many others.
The third way was opened by William Farr (1807-1883),
author of a "law" to the effect that "the curve of an epidemic at
first ascends rapidly, then slopes slowly to a maximum, to fall
more rapidly than it mounted." Greenwood, Brownlee, Ronald
Ross, and others have followed Farr and their epidemic curves
are usually of the normal bell-shaped Farr type. (13-26)
46,4 Russell: Avian Malaria Studies, III 653
Ross, in particular, (19-26) ever since his epochal discovery of
the transmission of malaria by mosquitoes, has been interested
in the mathematics of the spread of malaria. He has applied
the theory of probabilities to the statistical prognosis of epide-
mics, dealing particularly with the statics or equilibrium of
malaria. His equations have been carefully analyzed and ampli-
fied by Lotka, (27-29) who has dealt more especially with the
kinetics of malaria. Others who have dealt with malaria mathe-
matically are Waite(30) and McKendrich.(3i, 32) As yet, how-
ever, there are no data available for numerical comparison be-
tween mathematical formulae and observed conditions.
The second way of approach to the biological basis of epide-
mics— the way of direct experimental epidemiology — has only
recently been taken. Reports of L6ffler,(33) Danysz,(34,35)
Bahr,(36) Liston,(37) Xylander,(38) Muhlens,(39) and Bain-
bridge (40) were suggestive. The first direct attempt at experi-
mental epidemiology was made by Topley and his colleagues (41-
72) in England. They have carried out notable studies with
mouse pasteurellosis infections.
These men have studied the epidemiological history of a
"little community, wholly exempt from res angusta domi in any
sense of the phrase, well fed, well housed, with nothing to do
but eat, fight, make love, and sleep, shielded from contamination
by supermedical officers of health, and most efficient birth con-
trol."
They have "brought the doctrine of Epidemic Constitutions
within the compass of natural inquiry." (62)
One of their most important findings has been that "a pasteu-
rellosis will continue as a fatal infectious disease within a popu-
lation of mice replenished wholly by additions of normal animals,
not infected prior to immigration, over a period of more than
3£ years, that is through a period longer than a generation." (58)
In other words, the admission to a controlled community of mice
of individuals not having the disease that is epidemic in that
community is a danger to that community. Here then is a
suggestive lead for investigation in human herds, for much of
our community prophylaxis ignores completely incoming nor-
mal persons.
Outstanding and independent work in experimental epidemiol-
ogy in villages of mice has been done at the Rockefeller Institute
by Flexner, Webster, and associates. These studies have shown
that epidemics may arise from increased dosages of the patho-
654 The Philippine Journal of Science 1931
genie organism. There is acceleration or diminution in response
to factors that determine the susceptibility or resistance of the
population. It appears from this work that the course of events
in the epidemics was based on the distribution of the bacterial
parasite among the population at risk and the susceptibility of
the individuals comprising the population. Bacterial virulence
does not appear to have been a changing factor. Later studies
at the institute have been with respiratory infections in rabbits
and with fowl cholera. (73-123) Neufeld, Lange, and coworkers,
at the Robert Koch Institute in Berlin, have also investigated
problems of experimental mouse typhoid. (124-133)
Still other attempts to study epidemiology by controlled exper-
iments are those of Perla and Lurie (134-139) with artificially
induced epidemics of tuberculosis in rabbits. Koch (140) observed
that rabbits and guinea pigs exposed in the same room with
tuberculous animals for a longer period than four months not
infrequently acquire tuberculosis. Perla and Lurie have at-
tempted well-controlled experiments on the basis of observations
from spontaneous outbreaks.
There are many references in the literature to spontaneous
epidemics of disease among laboratory animals and these often
afford excellent although uncontrolled opportunities for study.
As an example the reports of Theobald Smith and Nelson, (141,
142) on paratyphoid in guinea pigs, may be cited.
Such natural episodes cannot, however, take the place of ex-
perimentally produced epidemics which, although simulating the
usual, are yet unusual in that certain factors are manipulated
and controlled in a uniform way.
These references to studies in the experimental epidemiology
of bacterial diseases are given not because they shed much light
on the spread of malaria in a community of birds. They are
cited, with such bibliography as is available to the author, be-
cause they illustrate a new method of approach to problems of
epidemiology, whether the disease in question be due to bacteria
directly passed from individual to individual or due to protozoa
carried by an arthropod host. Such experiments inaugurate a
new era in epidemiology.
It is undoubtedly open to question whether experimental epi-
demics within the confines of small cages accurately reflect the
phenomena of natural outbreaks of disease, and it is partly to
answer this question that the following experiments have been
46,4 Russell: Avian Malaria Studies, III 655
undertaken. While the immediate findings of the experiments
here reported are meager yet the general way of approach may
lead into fertile territory.
So far as the author is aware this is the first report to be
published on the experimental epidemiology of malaria, based
on miniature epizootics of the disease in laboratory animals.
There is a brief reference in Gill's excellent textbook on epide-
miology, to some abandoned studies with experimental malaria
in sparrows. No other has been found in the available litera-
ture.
GENERAL EPIDEMIOLOGY OP MALARIA
Certain fundamental considerations governing the spread of
malaria are well known. Before a new case can arise in a
community the following conditions must obtain progressively:
1. There must be a gametocyte carrier ; that is, a person with
sufficiently numerous and normal mature male and female game-
tocytes circulating in the peripheral blood, — the seed.
2. A female anopheline mosquito capable of acting as a bene-
ficent host to malaria parasites must travel to the skin of the
gametocyte carrier, push its proboscis into a blood vessel, and
suck enough gametocytes into its gut to insure that it will become
host to the critical number, or more, of sporozoites.
3. The mosquito that has thus successfully fed must live long
enough and must maintain conditions of temperature and bodily
state favorable enough to make possible the development of
malaria parasites to the sporozoite stage, with lodgement of these
sporozoites in the salivary glands.
4. This mosquito must make its way successfully to the skin
of a susceptible person and inject a sufficient number of sporo-
zoites to cause a new infection, — the sower and soil.
These fundamental and undisputed considerations remove
much of the mystery from malaria epidemics, but they have not
made the situation entirely clear. Anophelism sans malaria;
malaria sans anophelism ; years or regions of hyperendemicity ;
incidence regressions sans prophylaxis; recurrences coincident
with active control ; the effect of changed environment, of over-
crowding, of malnutrition; these and other phenomena of ma-
laria require further elucidation. The studies to which the
present paper is an introduction have been undertaken in the
hope that they may send light, however dim, into some of the
hidden recesses of the epidemiology of malaria.
656 The Philippine Journal of Science 1931
AVIAN MALARIA
The study of avian malaria has helped in the solution of some
of the problems of human malaria. MaeCallum's dicovery of
the exflagellation of Haemoproteus;(i^) Ross's momentous
discovery of the transmission of Proteosoma by mosquitoes; (145)
studies in relapse, drug therapy, biology and biometry of para-
sites, and host immunity in avian malaria by Whitmore,(i46)
the Sergents,(l47) Roehl, and others, (148) Hartman,(i49) and
Huff (150) have augmented our understanding of human malaria.
It seems logical, therefore, to expect that a careful experi-
mental study of epizootics of malaria in laboratory birds may
enrich our knowledge of the epidemiology of human malaria.
PROCEDURE
1. Birds. — The birds used in these experiments were canaries
(Serinus canarius), purchased from dealers. Up to the time
of writing this report two hundred canaries have been purchased
and examined, all being negative. In no case has a bird infected
with malaria been received from a dealer.
As will be seen below in the three preliminary reports of exper-
iments, there was a high mortality in the first case but a low one
in the second and third cases. This illustrates the fact that
while some lots of canaries do poorly under laboratory conditions,
others do very well.
2. Parasites. — The parasite used in all of these experiments
was Plasmodium cathemerium Hartman, 1927.(151) In the first
experiment the original Baltimore strain was used. In the last
two experiments a strain isolated by Huff in Boston was
used. (152) It is a matter of common knowledge among those
who have worked with this Plasmodium that canaries are suscep-
tible to it. In over two hundred cases, in the experience of the
author, it has invariably established itself in a bird upon needle
inoculation. Successful transmission by mosquitoes occurred in
the first and third experiments of this paper. It has also been
reported by Huff (153) and others.
3. Mosquitoes. — The mosquito used in the first experiment
was Culex (Culex) pipiens Linnaeus, 1758. In the second and
third experiments the species used was Culex (Culex) quinque-
fasciatus Say, 1823, (C. fatigans). That these species are sus-
ceptible to infection with avian malaria parasite has been shown
in general for C. quinquefasciatus by Ross, (145) Daniels, (154)
James, (155) and others. Huff (153) has shown in particular that
46, 4 Russell: Avian Malaria Studies, III 657
C. quinquefasciatus is susceptible to P. cathemerium Hartman,
1927. The susceptibility of C. pipiens has been demonstrated in
general by Huge, (156) the Sergeants, (157) and Neumann. (158)
Huff (153) has shown in particular that C. pipiens is susceptible
to P. cathemerium Hartman, 1927.
4. Environment— The cages have been approximately 3 by 2
by 2 feet in size (Plate 1) . In the first experiment the cage had
glass sides and was kept in a special room equipped with ther-
mostat and electric heater so adjusted that the temperature
remained at about 80° F. (range 79° to 82° F. or 26.0° to 27.7°
C). High relative humidity was maintained. The birds were
in a wire cage placed inside the mosquito cage in such a position
that cleaning and feeding could be carried on with a minimum
of disturbance.
The second and third experiments were done in Manila where
the temperature is at no time unsuitable for mosquito breeding.
Glass sides were not used in the Manila cages (Plate 2).
6. Controlled factors. — It will be realized from the foregoing
paragraphs that many factors underlying the spread of malaria
among birds could be manipulated in these experiments. A
definite number of susceptible individuals were shut in a con-
trolled area in close association with a definite number of game-
tocyte carriers. The carriers were changed from time to time,
as indicated by daily blood smears, in order to keep a plentiful
supply of gametocytes available.
Just how many gametocytes per 10,000 red blood cells are re-
quired to infect a mosquito is not known. It may be pointed
out that Darling(i59) found the limit of infectiousness in human
malaria from A. albimanus to be one gametocyte per 500
leucocytes or 12 gametocytes per cubic millimeter of blood. As
Huff (153) points out, if we assume the same to hold true for bird
malaria and C. quinquefasciatus and assume further an ansemia
of 2,500,000 red cells per cubic millimeter of blood, then a bird
with only 0.048 gametocyte per 10,000 red cells would be in-
fectious to mosquitoes.
In the experiments reported below, the carriers, with rare
exceptions, were found upon examination to have 5 or more
gametocytes per 10,000 red cells (see Tables 1, 2, and 3). If
we assume with Huff, following Darling's work again, that an
average blood meal is 0.0008 gram or 0.76 cubic millimeter (it
is probably more) and also assume again an anaemia of 2,500,000
red cells per cubic millimeter of blood (it is probably not so
658
The Philippine Journal of Science
1931
severe), then a single meal would mean 1,900,000 red cells and
in the present experiments, 950 or more gametocytes (often
several thousands).
Table 1. — First experiment. Gametocyte counts.
Date.
Bird No.
1929
April 17.
April 18_
April 19_
April 20.
April 21.
April 22.
April 23.
April 24.
April 25.
April 26.
April 27.
April 28.
April 29.
April 30.
May 1__
May 2.-
May 3._
May 4. _
May 5__
May 6. .
May 7__
2RH
2RH
2RH
115H
115H
115H
4RH
4RH
4RH
3RH
3RH
4RH
4RH
6RH
6RH
6RH
6RH
6RH
6RH
6RH
7RH
Smear.
Gameto-
cytes per
10,000
red blood
cells.
+ + + +
24
+ + +
31
+ + +
132
+ + + +
163
+ + + +
210
+ + +
92
+ + + +
20
+ + +
8
+ +
3
+ + +
47
+ + + +
108
+ + +
76
+ + +
55
+ + +
151
+ + + + +
132
+ + + + +
201
+ + + + +
154
+ + +
95
+ + + +
236
+ + + +
102
+ + + +
269
Date.
1930
May 8__
May 9__
May 10..
May 11 _.
May 12..
May 13..
May 14..
May 15.
May 16..
Mayl7_.
May 18..
May 19. .
May 20 _.
May21_.
May 22..
May 23. .
May 24 _.
May 25 _.
May26_.
May 27..
Bird No,
7RH
7RH
7RH
8RH
8RH
8RH
8RH
8RH
8RH
9RH
9RH
9RH
9RH
9RH
126H
126H
126H
43RE
43RE
43RE
Smear.
+ + + +
+ + + +
+ + + +
+ + +
+ + +
+ + +
+ + + +
+ + +
+ + + +
+ +
+ +
+ +
+ +
+ +
+ + +
+ + + +
+ + + +
+ + + + +
+ + + +
+ + + +
Gameto-
cytes per
10,000
red blood
cells.
730
619
988
50
32
49
83
76
51
21
17
28
7
5
113
420
519
581
690
834
Table 2. — Second experiment' Gametocyte counts.
Date.
1930
April 28.
April 30_
May 1__
May 4..
May 13.
May 14.
May 17.
May 26.
May 31.
June 4__
June 5__
Bird No.
21R
21R
21R
21R
48R
48R
48R
48R
52R
52R
51R
Smear.
Gameto-
cytes per
10,000
red blood
cells.
+ +
1
+ + + + +
114
+ + + + +
83
+ + + +
42
+ + + +
110
+ + +
155
+ +
95
+
5
+ + + +
34
+ + + + +
14
+ + + + +
317
Date.
Bird No.
1930
June 9 _ _ .
June 10 _.
June 21 _.
June 24 _.
July4.._.
July7.._.
July 10__.
July 13...
July 15...
July 18...
July 24.. .
51R
43R
43R
33R
33R
34R
34R
U31
U30
U30
U2
Smear.
Gameto-
cytes per
10,000
red blood
cells.
+ +
+ + +
+ + + + +
+
+
0
+
+ + + +
+ + + + +
+ + + + +
21
31
1,070
18
5
1
0
6
25
257
209
46,4 Russell: Avian Malaria Studies, HI 659
Table 3. — Third experiment. Gametocyte counts.
Date.
Bird No.
Smear.
Gameto-
cytes per
10,000
red blood
cells.
Date.
Bird No.
Smear.
Gameto-
cytes per
10,000
red blood
cells.
1930
1930
August 14
U40
+ + + +
310
August 30
U73
+ + + + +
891
August 14
U41
+ +
35
August 30
U22
+ + + +
364
August 14
U42
+ + +
29
August 30
U38
+ + + + +
403
August 14
TJ36
+ + +
47
September 8 _ _
U84
+ + + + +
905
August 21
U36
+ + +
54
September 8__
U38
+
2
August 21
U88
+ + + +
286
September 8__
U93
+ + + +
171
August 21
U89
+ + +
71
September 8__
X12
+ + + + +
238
August 21
U91
+ + +
92
September 16 _
U80
+ +
23
August 30
U37
+ + + + +
1,037
September 16 _
J38
+ + + + +
1,001
The mosquitoes were grown by the usual technic, for the most
part inside the cage itself where feeding, mating, and egg-laying
proceeded without difficulty. Daily counts were made of the
adult mosquitoes, and from time to time increments of larvae and
pupae were added to the cages (see Tables 5 and 6). Loffler's
dehydrated blood serum mixed with litmus milk was found to
be a satisfactory food for the larvae. Raisins in sugar syrup
were supplied for the male adult mosquitoes.
As to the susceptibility of the mosquitoes used in the first
experiment it may be placed at about 45 per cent in accordance
with results given in Table 4 of the very complete paper by
Huff. (153) I was indebted to Huffi for the stock of C. pipiens,
which was the same as used in his experiments. At this point
I would state that I am indebted to him not only for mosquito
stock but also for generous criticism and guidance in many
phases of this work.
In the second two experiments the susceptibility of the stock
of C. quinquefasciatus may be placed for the purposes of this
paper at about 48 per cent. This is based on one experiment
where females of this species were allowed to feed on gametocyte-
carrying birds so placed as to allow the insects leisurely and
complete meals. Of thirty-one individuals that survived twelve
days, fifteen on dissection were found to have oocysts on the wall
of the mid-gut. Experiments are in progress to determine the
sporozoite rate which is lower.
As to the biting-frequency factor, data are being gathered
but are not yet available.
660
The Philippine Journal of Science
1931
As to the longevity of the mosquitoes in the experimental
cages, Tables 4, 5, 6, and 7 give some preliminary information.
Further observations are in progress.
Table 4. — Experiment 5, cage E-H. Mortality of mosquitoes in an expert-
ment cage (C quinquefasciatus).*
Date.
Adults
counted.
Pupae
added.
Egg rafts
removed.
Date.
Adults
counted.
Pupae
added.
Egg rafts
removed.
1931
January 20_
176
63
160
88
138
154
74
60
42
19
8
17
1931
February 11
February 12
February 13
February 14
February 15
February 16
February 17
February 18
February 19
February 20
February 21
February 22
February 23
February 24
February 25
February 26
February 27
February 28
March 1
37
36
26
26
10
10
9
9
9
9
5
5
5
5
5
5
2
2
1
0
0
0
January 21
January 22
January 23 _.
January 24
January 25
January 26
January 27 __
January 28
January 29
January 30
January 31
February 1
February 2
February 3
February 4
February 5
February 6
February 7
February 8
February 9
February 10
73
100
135
192
212
295
250
280
250
237
137
68
110
100
37
37
36
39
39
39
39
1
4
2
2
1
March 2
March 3
March 4
a Throughout this experiment two birds were kept in the cage to supply blood meals.
Raisins in syrup were supplied for the males.
FIRST EXPERIMENT
Chart 1 illustrates the following summary of experiment 1.
February 17, 1929. Three hundred fifty larvaa and pupae of C. pipiens
put in special cage with nine canaries (none infected) ; temperature, 85°
F.; humidity, 95 per cent.
March 1. First egg raft found inside cage.
April 17. Mosquitoes in cage feeding, mating, laying eggs, and dying at
such a rate that the daily count of females averages 100. Susceptible
birds 1 to 9R put in cage. Also gametocyte carrier 2RH.
April 20. Gametocyte carrier 115H substituted for 2RH.
April 22. Susceptible 3R died with no evidence of malaria. 10R put
in cage as replacement.
April 23. Gametocyte carrier 4RH substituted for 115H.
April 26. Gametocyte carrier 3RH substituted for 4RH.
April 28. Gametocyte carrier 4RH substituted for 3RH.
46,4 Russell: Avian Malaria Studies, HI 661
Table 5. — Mosquito population during second experiment.*
Date.
Mosqui-
toes
counted.
Date.
Mosqui-
toes
counted.
405
415
421
378
271
372
307
343
408
478
435
456
557
576
604
621
846
812
880
746
669
663
692
677
656
683
669
783
718
727
Date.
Mosqui-
toes
counted.
1930
May 16
758
877
894
888
829
815
798
749
817
791
644
569
442
338
310
312
319
382
383
364
456
457
496
503
403
440
405
459
420
413
1930
June 15
1930
July 15
635
639
653
635
651
496
571
492
516
543
535
454
418
445
496
530
513
460
458
456
457
432
429
421
412
438
441
482
439
425
May 17
June 16 _
July 16__
Mayl8_
June 17.
July 17. .
May 19
June 18
June 19
July 18
May 20
July 19
May 21 _.
June 20 . _
July 20 _
May 22
June21___
July 21
May 23
June 22
July 22
May 24
June 23 __
July 23
May 25
June 24
July 24
May 26
June 25
July 25
May 27 _
June 26
July 26
May 28
June 27 _ _ _ _
July 27
May 29
June 28
July 28
May 30
June 29
July29_
May 31
June 30
July 30
June 1
Juiyl
July 31
June 2
July 2
August 1
June 3
July 3__
August 2
June 4 ._
July 4
August 3
June 5
July 5
August 4
June 6
July 6. _
August 5
June 7
July 7
August 6
June 8
July 8
August 7
June 9
July 9
August 8
June 10 ___
July 10__
August 9
June 11
July 11„ .
August 10
June 12 __
July 12
August 11
June 13 .
July 13
August 12
June 14 _ _
July 14
August 19
aIn this experiment the egg rafts were not removed from the cage. Additional pupse
were added from time to time as the population seemed to be falling. All counts Were
made at about 9 a. m. almost always by the same two individuals, their totals being aver-
aged.
April 30. Gametocyte carrier 6RH substituted for 4RH.
May 3. Susceptible 1R died with no evidence of malaria; replaced by
11R.
May 4. Susceptible 4R died with no evidence of malaria; replaced by
12R.
May 5. If a maximum time of twelve days is allowed for development
of sporozoites and six days for a prepatent period in a new infection, a
case of malaria would appear on this day had a mosquito become infected
the first night and lived to bite another bird on the twelfth day.
May 7. Gametocyte carrier 7RH substituted for 6RH. Susceptible
10R died with no evidence of malaria; replaced by 13R.
May 8. Susceptible 8R died with no evidence of malaria; replaced by
14R.
662 The Philippine Journal of Science
Table 6. — Mosquito population during third experiment*
1931
Date.
Mosqui-
toes
counted.
Date.
Mosqui-
toes
counted.
Date.
Mosqui-
toes
counted.
1930
August 15
806
881
859
847
779
757
723
670
390
423
518
662
548
1930
August 28
557
459
629
792
1,043
875
735
631
639
812
821
814
1930
September 9_
September 10
September 11
September 12
September 13 _
September 14
September 15
September 16
September 17
September 18 . _
September 19
September 20
834
949
896
1,061
863
834
849
961
863
743
705
717
August 16.^
August 29
August 17
August 30
August 18
August 31 _
August 19
September 1
September 2 _ _
September 3
September 4
September 5
September 6
September 7
September 8
August 20
August 21
August 22
August 23.
August 24
August 25
August 26
August 27
* In this experiment egg rafts were removed from the cage (see Table 7). Pupae were
added from time to time as the population seemed to be falling. The counts were made
chiefly by one individual and usually at 9. a. m.
Table 7. — Egg rafts removed from cage during third experiment.
Date.
Egg
rafts
remov-
ed.
Date.
Egg
rafts
remov-
ed.
Date.
Egg
rafts
remov-
ed.
Date.
Egg
rafts
remov-
ed.
1930
1930
1930
1930
August 19
5
August 30
47
September 14 __
6
September 23 __
2
August 20
5
August 31
52
September 15 __
5
September 24. _
36
August 21
4
September 3___
10
September 16 __
5
September 25. .
23
August 22
15
September 5
3
September 17 __
2
September 26 __
0
August 23
1
September 6___
106
September 18 __
32
September 27 __
0
August 24
6
September 8
22
September 19. _
64
September 28. _
28
August 25
3
September 9 ...
66
September 20..
6
August 26
3
September 10 __
10
September 21 __
12
August 28
7
September 11 __
36
September 22 __
2
May 11. Gametocyte carrier 8RH substituted for 7RH. Susceptible 11R
died with no evidence of malaria; replaced by 15R.
May 14. Susceptible 2R and 13R died with no evidence of malaria;
replaced by 16R and 17R.
May 17. Gametocyte carrier 9RH substituted for 8RH.
MJay 22. Gametocyte carrier 126H substituted for 9RH.
May 23. Susceptible 6R and 16R died with no evidence of malaria;
replaced by 11RH and 13RH.
May 25. Gametocyte carrier 43RE substituted for 126H. Susceptible
5R and 15R died with no evidence of malaria; not replaced1 because birds
not available.
46,4 Russell: Avian Malaria Studies, III 663
May 27. Replacements 37RE and 40RE put in cage. This day is not-
able for the fact that both 12R and 14R have positive blood smears.
May 29. Susceptible 17R died with no evidence of malaria; not re-
placed.
May 30. Replacement 45RE.
June 4. Bird 12R died of acute malaria.
June 5. Bird 9R has a positive blood smear. It has been in the cage
since April 17 and escaped infection from about April 29 to about May 30.
May 6. Bird 14R died of acute malaria.
May 7. Bird 40E died with no evidence of malaria.
May 9. Bird 45RE died with no evidence of malaria.
May 11. Experiment stopped.
DISCUSSION OF FIRST EXPERIMENT
This first experiment was carried through to develop technic.
It demonstrated that C. pipiens will propagate itself and main-
tain a colony in an experimental cage such as described. It
made evident that C. pipiens feeds readily on birds in cages and
that it will transmit malaria under these conditions. It also
demonstrated that a high mortality may come about among
birds free from malaria but purchased in the open market and
kept under laboratory conditions. Each dead bird was carefully
examined post mortem for evidence of malaria. Except in the
cases of 12R and 14R no such evidence was found. The other
birds died of a bacterial infection. This bacterial epidemic
coincident with the experiment illustrates one of the occasional
major difficulties in the study of avian malaria.
SECOND EXPERIMENT
Chart 2 illustrates the second experiment. Having moved
from Boston to Manila it became necessary for me to develop a
technic suited to different climatic conditions. This second ex-
periment and the third are to be viewed in a preliminary way.
April 5, 1930. Four pans well stocked with larvae and pupae of C.
quinquefasciattts put in cage with five birds.
April 12. First egg rafts.
April 27. Colony of mosquitoes is propagating itself strongly.
April 28. Susceptible birds XI, 2, 3, 4, 5, 6, 8, 9, and 10 put in cage.
Also gametocyte carrier 21R.
May 13. Gametocyte carrier 48R substituted for 21R.
May 27. Gametocyte carrier 52R substituted for 48R.
June 4. Gametocyte carrier 51 R substituted for 52R.
June 10. Gametocyte carrier 43R substituted for 51R.
June 21. Gametocyte carrier 33R substituted for 43R.
July 7. Gametocyte carrier 34R substituted for 33R.
July 11. Gametocyte carrier U31 substituted for 34R.
664
The Phttippine Journal of Science
1931
APRIL i
t
CHART
t «
NUMBER
J « 7
» FIRST
• » (0 11 15
EXPERIMENT
13 H It IS
EPIDEMIOLOGY OF AVIAN
IT IS IS JO Jl n 23 24 « S«
MALARIA
27 28 29
SO 31
IR
O
o
0
2R
0
o
o
3R
0 D
4R
0
o
o
5R
O
o
o
6R
o
o
o
7R
o
o
o
6R
o
o
0
OR
o
0
o
fOR
, 1°
o
o
2RH
♦+♦♦ -m ++4-J
II5H
|+4-4-»-4f4-4 4-4*
4RH
.444+444
+4
4-4+ 44 +
3RH
+4 +
♦ «♦♦
6RH
MAY
IR
D
2R
0
o
O
OD
4R
0
D
5R
0
0
O
0
0 0 o
0
6R
0
0
0
0
0 0 OD
7R
0
0
o
0
0 0
0
0
8R
0
0
OD
9R
o
0
0
0
0 o
0
0
IOR
o
00
IIR
°
o
OD
I2R
0
0
O
0
0 o
+
++
I3R
0
0
OD
I4R
°
0
0
O 0
+
4 +
I5R
o
0
o 0
D
I6R
O O
O O OD
I7R
0 O
0 0
o
D
MRH
o
0
0
I3RH
0
0
0
37RE
0
0
dORE
0
0
45RE
1
0
6RH *M
,.-.♦
♦♦♦
♦-♦♦♦«
7RH
m*
M44f4+4 *»♦
8RH
tH ♦♦♦• ♦ ♦+
*♦♦» ***t+*+
9RH
t+ ♦ ♦ +♦ +t 4-4-
I26H
^
\
43RE
mn nt
♦ tf
JUNE
7R
0
o
o
°l
9R
0
++
I2R
+
D
I4R
++
0
IIRH
0
o
o
o
I3RH
0
o
o
0
37RE
0
o
0
0
dORE
0
o
D
45RE
0
0
0 D
-
Fig. 1. Chart 1, first experiment, epidemiology of avian malaria.
46'4 Russell: Avian Malaria Studies, HI 665
July 14. Gametocyte carrier U30 substituted for U31.
July 22. Gametocyte carrier U2 substituted for U30.
July 24. X8 died with no evidence of malaria. This is the first bird
to die in the cage since the beginning of the experiment. Gametocyte
carrier U2 removed and not replaced.
July 28. X9 died with no evidence of malaria.
August 7. X2 died with no evidence of malaria.
August 12. Experiment discontinued. There has been no transmis-
sion of malaria whatever in this experiment, yet at all times there have
been gametocytes available.
DISCUSSION OF SECOND EXPERIMENT
In order to test the susceptibility of the birds in this experi-
ment XI, 3, 4, 5, 6, and 10 were inoculated by needle with the
same strain of malaria Plasmodium on dates as shown in fig. 2.
Birds X3 and X10 died before becoming positive, four and two
days, respectively, after inoculation. The other four birds be-
came positive after the usual prepatent periods, showing that
they were susceptible and proving that they had not previously
been infected by the mosquitoes.
The same strain of mosquitoes was used in the third experi-
ment (see below) and they were thereby also proved to be sus-
ceptible.
In this second experiment we therefore had a situation cor-
responding somewhat to anophelism sans malaria. The mosqui-
toes readily fed on the population and were not diverted to other
animals, as is sometimes the case in nature where anophelism
sans malaria exists. At all times 10 per cent (one bird) of the
population carried gametocytes, yet malaria did not spread.
There was too high mortality among the mosquitoes and there
were too few gametocyte carriers. The mosquito turnover was
much higher in this experiment than in the first.
The "epidemic potential" in this second experiment was not
high enough. This term, "epidemic potential," was suggested
by Peters, (160) whose book is not available to this author.
Topley defines Peters's term as "the balance of interacting forces
which tends towards the occurrence of an outbreak of disease."
It is a good term to replace Sydenham's ill-defined "epidemic
constitution," which seems to have been forced back into use by
the last pandemic of influenza, an occurrence that was and is
inexplicable in terms of modern epidemiology. "Epidemic po-
tential," as a term, helps not at all toward fundamental explan-
ations, yet it more aptly expresses the modern view.
264209 9
666
The Philippine Journal of Science
1931
CHART NUMBES
StCONO EXPERIMENT EPIDEMIOLOGY OF AVIAN MALARIA
52R _
JUNE
51 R
HP.
33A
JULY
33* i
3JR
0
0
0
IMS
♦ ♦♦♦ ♦♦
♦
0
p
0
0
0
0 1/73
D
0
0
0
o im
0
0
0
o
ltfl
o
0
0
0
m»i
0
0
0
0 WJ7
0
Fig. 2. Chart 2, second experiment, epidemiology of avian malaria.
46,4 Russell: Avian Malaria Studies, HI 667
THIRD EXPERIMENT
A colony of C. quinquefasciatus having established itself in a
cage, the experiment was started.
August 14, 1930. Susceptible bird X13 put in cage with gametocyte
carriers U36, U40, U41, and U42.
August 19. Gametocyte carriers U48 and' U88 substituted for U41 and
U42.
August 21. Gametocyte carriers U89 and U91 substituted for U40
and U48.
August 23. Gametocyte carrier U90 substituted for U36.
August 27. Gametocyte carriers U37 and U73 substituted for U88
and U89.
August 28. Gametocyte carriers U90 and U91 removed from cage.
August 29. Gametocyte carriers U22 and U38 put in cage.
September 2. Gametocyte carriers U68, U69, U72 substituted for U22.
U37 and U73.
September 5. Gametocyte carriers U92, U93, and X12 substituted for
U68, U69, and U92.
September 6. Gametocyte carrier U92 removed from cage.
September 8. Gametocyte carrier U84 put in cage.
September 9. Gametocyte carriers U78 and J17 substituted for U38
and X12.
September 11. Gametocyte carrier J18 substituted for U93.
September 12. Gametocyte carrier J22 substituted for U84.
September 13. Gametocyte carrier J40 substituted' for J22.
September 15. Gametocyte carriers U80 anad J38 substituted for U78
and J40.
September 17. XI 3 has a positive blood smear. Experiment ended.
DISCUSSION OF THIRD EXPERIMENT
Here we had a condition simulating an area in which malaria
is hyperendemic and into which a susceptible individual comes.
Eighty per cent of the population carried gametocytes in their
blood. There were mosquitoes in abundance. It was a foregone
conclusion that the susceptible would become infected.
The mosquito turnover in this third experiment was compar-
able to that in the second where no transmission of malaria
took place. But here the "epidemic potential" was high.
These first experiments are not to be analyzed too closely
mathematically. It has been necessary to proceed slowly and
to develop technical facility. This report is made as an intro-
ductory one because it seems to point towards an experimental
procedure in the study of the epidemiology of malaria which
may lead to a better understanding of fundamental relationships.
668
The Philippine Journal of Science
1931
AUGUST
XI3
U36
U40
U4I
U42
U48
U88
U89
U9I
U»0
U37
U73
U22
U38
SEPT.
XI3
U37 <
U73 •
U22
U38 *****
U68
U69
U72
U92
U93
XI2
U84
U78
JI7
J 16
J 22
J40
U80
J38
GM*«T W/M6ER III THIRD EXPERIMENT EPIDEMIOLOGY OF AVIAN MALARIA
j a * »• i » 8 10 11 12
1* 16 1« II II
O
+♦♦♦♦»♦♦♦*+♦♦ +♦ ♦+ ♦♦♦ ♦♦+
♦♦♦♦ ■»♦ ♦ + ♦+ + I
♦•+♦ ♦-♦ 4-
*» M » «» f« XI f* t* 49 91
o a
***■ ♦+* ♦♦♦+ ++♦ ♦•♦
h ♦*+♦•*♦»» t *»+♦»*«»♦*»
0 0 0 0
Pig. 8. Chart 3, third experiment, epidemiology of avian malaria.
SUMMARY
The subject of experimental epidemiology is briefly discussed
with especial reference to malaria. Three preliminary experi-
ments in the epidemiology of avian malaria are reported with
a discussion of technic and results. A bibliography is appended.
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132. Okamoto, T. Epidemiologische Beobachtungen an Mausen und Neer-
schweinchen. Klin. Wochenschr. 5 (April, 1926) 795-796. Japan
Med. World 6 (August, 1926) 210-213. Abs. Journ. Am. Med. Assoc.
86 (June, 1926) 2006.
676 The Philippine Journal of Science im
133. Neufeld, F. Natural immunity in its significance for epidemiology.
De Lamar Lectures (1926-1927) 1-12.
134. Perla, D. Experimental epidemiology of tuberculosis. Journ. Exp.
Med. 45 (1927) 209-226.
135. Perla, D. Experimental epidemiology of tuberculosis. The elimina-
tion of tubercle bacilli in the feces, bile and urine of infected guinea
pigs. Journ. Exp. Med. 45 (1927) 1025-1036.
136. Lurie, M. B. Experimental epidemiology of tuberculosis. The effect
of crowding upon tuberculosis in guinea pigs, acquired by contact
and by inoculation. Journ. Exp. Med. 51 (1930) 729-741.
137. Lurie, M. B. Experimental epidemiology of tuberculosis. Air-borne
contagion of tuberculosis in an animal room. Journ. Exp. Med. 51
(1930) 743-751.
138. Lurie, M. B. Experimental epidemiology of tuberculosis. The route
of infection in naturally acquired tuberculosis of the guinea pig.
Journ. Exp. Med. 51 (1930) 769-775.
139. Lurie, M. B. Experimental epidemiology of tuberculosis. The effect
of eliminating exposure to enteric infection on the incidence and
course of tuberculosis acquired by normal guinea pigs confined with
tuberculous cage mates. Journ. Exp. Med. 51 (1930) 753-767.
140. Koch, R. Gesammelte Werke von R. Koch, Leipsic 1 (1912) 512.
141. Nelson, John B., and Theobald Smith. Studies on a paratyphoid
infection in guinea pigs. I. Report of a natural outbreak of para-
typhoid in a guinea pig population. Journ. Exp. Med. 45 (1927)
353-364.
142. Smith, Theobald, and J. B. Nelson. Studies on a paratyphoid infec-
tion in guinea pigs. II. Factors involved in the transition from
epidemic to endemic phase. Journ. Exp. Med. 45 (1927) 365-378.
143. Gill, C. A. The Genesis of Epidemics. Wm. Wood & Co. New York
(1928).
144. MacCallum, W. G. On the haemetozoan infection of birds. Journ.
Exp. Med. 3 (1898) 117-136.
145. Ross, R. Report on the cultivation of Proteosoma Labbe in the grey
mosquito. Calcutta. Reprinted in the Ind. Med. Gaz. 33 (1898) 410.
146. Whitmore, E. R. Observations on bird malaria and the pathogenesis
of relapse in human malaria. Johns Hopkins Hosp. Bull. 29 (1918)
325.
147. Sergent, Ed. and Et. A vantages de la quininisation preventive
demontr^s et precises experimentalement (paludisme des oiseaux).
Ann. Inst. Past. 35 (1921) 125-141.
148. Roehl, W. Die Wirkung des Plasmochins auf die Vogelmalaria.
Arch. f. Schiffs- u. Trop.-Hyg. 30 (1926) 11-18.
149. Hartman, E. Certain interrelations between P. praecox and its host.
Am. Journ. Hyg. 7 (July, 1927) 407-432.
150. Huff, C. G. The effects of selection upon susceptibility to bird ma-
laria in Culex pipiens Linn. Ann. Trop. Med. & Parasit. No. 4 23
(December 31, 1929).
151. Hartman, E. Three species of bird malaria, P. praecox, P. catheme-
rium sp. no v., P. inconstans sp. no v. Archiv f. Protistekunde 60 (1)
(December 2, 1927) 1-7.
4«,4 Russell: Avian Malaria Studies, HI 677
152. Huff, C. G. Personal communications.
153. Huff, C. G. Studies on the infectivity of Plasmodia of birds for mos-
quitoes with special reference to the problem of immunity in the
mosquito. Am. Journ. Hyg. No. 6 7 (November, 1927) 706-734.
154. Daniels, C. W. On the transmission of Proteosoma to birds by the
mosquito. Roy. Soc. Rep. to the Mai. Com. Proc. Roy. Soc. London
44 (1899) 443-454.
155. James, S. P. Malaria in India. Sc. Mem. by the Off. of the Med.
Journ. San. Depts. of Gov. of India. New Series No. 2 (1902).
156. Ruge, R. Untersuchungen uber das deutsche Proteosoma. Centralbl.
f. Bakt. 29 (1901) 187-191.
157. Sergent, Ed. and Et. Les hemalozoaires d'oiseaux. Ann. PInst. Past.
21 (1907) 251.
158. Neumann, R. 0. Die Ubertragung von Plasmodium praecox auf Kana-
rien vogel. Arch. f. Protist. 13 (1909) 23-69.
159. Darling, S. T. Factors in the Transmission and prevention of mala-
ria in the Canal Zone. Ann. Trop. Med. & Parasit. 4 (1910) 179-225.
160. Peters, O. H. Epidemic Diarrhoea. Cambridge University Press
(1911).
ILLUSTRATIONS
Plate 1
Cage used in first experiment in Boston. (The front glass has been
raised.)
Plate 2
Fig. 1. Type of cage used in Manila; front view.
2. Type of cage used in Manila ; end view.
TEXT FIGURES
Fig. 1. Chart 1, first experiment, epidemiology of avian malaria.
2. Chart 2, second experiment, epidemiology of avian malaria.
3. Chart 3, third experiment, epidemiology of avian malaria.
679
Ri'.ssM.r,: Aman Mvi.\iu,\ Sti-kiks, IIL|
ll'i
•• /
PLATE 1.
Russell: Avian II
I*. N<
*™2
Wi
mi
* s^M£
v*;-m§
PLATE 2.
BORED-HOLE LATRINE EQUIPMENT AND
CONSTRUCTION *
By Clakk H. Yeager
Of the International Health Division, Rockefeller Foundation
SEVEN PLATES AND FORTY-SIX TEXT FIGURES
In response to numerous requests for descriptions of latrine-
boring equipment, where it may be purchased, the cost, and how
to use it, this article has been prepared.
The selection of the locations for bored-hole latrines should be
under the direction of a person who has been instructed as to
the possibility of pollution of domestic water supplies, especially
the contamination of nearby shallow wells. Until more scienti-
fic data are available, the installation of bored latrines is sug-
gested only for areas in which there is no danger of infecting
the drinking water.
Satisfactory boring equipment that is cheap enough for wide
distribution in poor communities is not yet made by any of the
manufacturers, but an inexpensive satisfactory auger can be
made by purchasing an auger bottom, and making the shaft and
turning handle locally.
According to requests there seems to be need for an auger
costing about 20 dollars United States currency that can be
used universally and will work in water, soft sand, mud, clay,
sea shells, ashes, and rock. We have not yet found a cheap
auger that will work everywhere, and it is not likely that a
single cheap auger can be made that will work in all the dif-
ferent formations. However, to be of practical value in most
places the boring equipment must be the cheapest that will do
the work. Many of the elaborate machine-driven rotary drills
used in the oil fields and in some mining operations are too ex-
pensive and are not practical for boring latrines. In many
places 90 per cent of the boring is in sandy clay or similar
material, and only 10 per cent in soft sand requiring valve tools
lrThis work was done with the support and under the auspices of the
Government of the Philippine Islands and the International Health Divi-
sion of the Rockefeller Foundation.
264209 — 10 681
682 The Philippine Journal of Science 1931
or in other formations in which more-expensive equipment is
needed. In such places it is a waste of money to purchase
the more-expensive augers for the entire area, when cheaper
augers will do 90 per cent of the work just as well and frequently
better because of the design of the auger.
In a previous article an auger was suggested that will bore
satisfactorily in most places, but this auger was not designed to
cut through rock or to work in quicksand. Complaints stating
that the auger failed in laterite, adobe rock, and sand have been
received. We would not select an auger made for wood boring
to cut a hole in marble or steel, and the selection of an auger is
just as important in earth boring. The selection of boring tools
depends upon the geological formation into which the hole is
to be made.
Much of the equipment to be described can be made locally.
It would require several volumes to cover the manufacturers'
descriptions of the boring equipment on the market. I have not
seen all of the augers manufactured, and very likely there are
good tools that have not come to my attention, but the ma-
terial included in this article will be sufficient for a selection of
supplies that will be useful in making bored-hole latrines in a
variety of formations. Much description that would not add
greatly to the value of the article has been omitted.
The size of the holes to be bored is important in selecting
equipment. Bored-hole latrines have been made from 10 to 24
inches in diameter, but 14- and 16-inch holes have been most fre-
quently bored. There are some advantages in boring the holes
only 12 inches in diameter, although there is more danger from
fouling the sides and the capacity is more limited. It has not
definitely been determined whether or not the 12-inch holes
give the soil bacteria a better chance to work on the contents
than in the larger holes. If the soil is porous the 12-inch holes
could be used several years, especially with the water trap dis-
cussed under Construction.
If the walls of the hole cave in, a lining is required and this
reduces the diameter of the 12-inch holes to about 10 inches,
which is too small for practical use. An advantage of the 12-
inch hole is that it is easier to bore than a 16-inch hole, and
lighter and less-expensive equipment can be used. In some
areas in the Philippines the 16-inch holes have been filled two-
thirds full within fifteen months because coconut husks, sticks,
and leaves were thrown into the hole instead of water or paper.
Twelve-inch holes in these places would be filled in less than
Y eager: Bored-hole Latrines
683
a year. In many countries 14-inch augers are used, but fouling
has been reported and in some of these places they have now
changed to 15- or 16-inch augers. However, the 14-inch holes
are satisfactory in many countries, and in the Philippines we
have installed many 14-inch latrines. If the soil is likely to
cave in we usually use a 16-inch auger because the bamboo lin-
ing reduces the diameter of the latrine to about 14 inches.
In letters from manufacturers larger holes have been recom-
mended in order to allow a man to go down to remove stones
when necessary, but the employees in the Straits Settlements,
Philippine Islands, and other countries have no trouble in going
down a 16-inch hole. Fortunately bowlders are seldom encoun-
tered, except in some areas.
A 14-inch latrine would have to be bored to a depth of about
26 feet to have the same capacity as a 16-inch latrine 20 feet
deep.
We generally speak of boring 14- or 16-inch holes with augers
of these diameters, but in most soils the augers actually cut
the holes an inch or two larger in diameter than the size of the
auger.
The depth of the bored-hole latrine is from 12 to 26 feet. We
stop at 12 feet only when deeper
boring is difficult and expensive.
The holes should average about
19 feet in depth, although we
have made a number of school
and public latrines 23 feet deep.
THE IWAN POST-HOLE AUGER
The I wan post-hole auger (fig.
1) is inexpensive and can be
used for nearly all surface-soil
borings where hardpan, rock,
and quicksand are not encount-
ered. The shaft of this auger
supplied by the manufacturer is
not as satisfactory as the shafts
made locally; so we purchase
only the "auger bottoms," as
the manufacturer calls them, and make our own shafts. The
auger bottoms consist of two cutting blades and an arch or
Fig. 1. The Iwan auger, a, Shaft attached
to auger arch ; b, a more solid joint
with a nut below the socket. The bolts
and nuts are not necessary if the shaft
is welded to the arch.
684 The Philippine Journal of Science iosi
yoke to which the blades are riveted and to which the shaft
can be attached. In order to save cost of transportation one
shipment of auger blades and arches were ordered unassembled.
We had the blades and arches welded and riveted together local-
ly and the augers worked very well, but we did not save much
money and had the additional trouble and loss of time. In
following orders we always ordered the "auger bottoms" as ad-
vised by the manufacturer and not the blades and arches. The
blades must be riveted in proper alignment or the auger will
not bore a straight hole. Because of easier packing in standard
cases we now order in lots of even dozens and not ten, six-
teen, or thirty-eight auger bottoms for one shipment. The bot-
toms 16 inches in diameter cost 81.60 dollars United States
currency a dozen. The 14-inch bottoms cost 67.20 dollars a
dozen.
We have broken only two of the 1/8-inch blades on the stand-
ard Iwan augers in the Philippines; therefore, for general dis-
tribution we do not find the additional cost of heavier blades
justified, but Dr. John F. Kendrick, in India, has not been so
fortunate, and purchases especially heavy blades and makes the
arches (yoke pieces) and shafts locally. Unusual soils require
the heavy blades. Iwan Brothers, of South Bend, Indiana,
make extra heavy 16-inch diameter auger blades of 3/16-inch
steel in lots of ten pairs at 6 dollars United States currency
per pair, f . o. b. cars, New York. The steel strap to bring the
blade points together at the bottom is furnished by the manu-
facturer. The auger bottom, complete with arch and heavy
blades riveted together, costs 12 dollars. The standard weight
16-inch augers cost 7.50 dollars each. The regular 14-inch au-
gers cost 5.95 dollars each, and according to Mr. Rollin C. Dean,
of the Rockefeller Foundation, the 14-inch augers with 3/16-
inch blades would cost about 10.50 dollars each.
The shaft. — The 1-inch shaft supplied by the manufacturers
of the Iwan augers is in short sections and has threaded joints.
This type of shaft is satisfactory for the purpose originally
intended as a post-hole auger, but for boring to a depth of about
19 feet a l^-inch shaft from 19 to 22 feet long is more satis-
factory.
The auger bottom measures 19 inches from the tip of the
blades to the top of the arch; therefore, a 20-foot shaft allows
boring to a depth of about 20 feet. We usually use ordinary
46> 4 Y eager: Bored-hole Latrines 685
water or gas pipe, plain or galvanized, which is shipped in
lengths averaging over 19 feet. The longest lengths are se-
lected because the auger is more easily turned if at least 2
feet of the shaft extends above the surface of the earth when
the last few turns are being made.
In soft soil li-inch pipe (inside diameter) for making shafts
has given satisfactory service, but in most of the soil in the
Philippine Islands it has been necessary to use l-|-mch pipe,
and in one place in the Straits Settlements 2-inch steam pipe
was used. In early trials the 1-inch and lf-inch shafts bent
and broke, and a number of braces designed to keep the shaft
from bending and boring at an angle were tried. These de-
vices worked very well, but added to the expense, complicated
the equipment, and required more time on account of lost mo-
tion in adjusting the shaft. Then shafts of larger diameter
were used, and by selecting shafts large enough to suit the soil
there was no further difficulty. Shafts that do not bend too
much will bore straight holes unless a bowlder or other solid
material causes deviation. A large shaft has a more even torque
than a small one and will not break or bend enough to bore
at an angle. Two- or 3-inch shafts can be used, but in most
hard soil l-|-inch pipe serves the purpose.
In places where the ordinary l|-inch water-pipe shafts broke,
steam pipe and boiler tubes were tried and not only extra
thick but double extra heavy pipe was used, but these
heavy pipes are usually not as satisfactory, considering cost and
service rendered, as ordinary pipe of larger diameter. The
strength for the money expended up to certain limits depends
upon the diameter of the shaft. The torque of an ordinary 1|-
inch water-pipe shaft is very much better than that of a solid
bar or a smaller pipe of the same length and weight. In some
countries the water pipe is of inferior quality and at times
crushes or splits up the seam; in these instances, after the
failure of 2-, 2|-, or 3-inch shafts, it might be necessary to
use heavy or extra-heavy steam pipe.
The shaft auger-bottom joint. — The threaded shafts of the
Iwan augers as supplied by the manufacturer are screwed into
a socket in the auger arch. These are probably satisfactory
for boring shallow holes for telegraph poles, but are not strong
enough for most bored-hole latrine boring. A more satisfactory
joint is shown in fig. 1, a. This joint is made by threading the
686 The Philippine Journal of Science 1931
end of a piece of round bar steel and screwing the bar into the
auger socket. A hole is drilled through the socket and bar and a
tool-steel pin is inserted for additional strength. A bar 10 inches
long and 1^ inches in diameter is suitable for a l|-inch shaft.
The shaft is fastened to the bar with two f-inch bolts or steel
rivets, which are inserted through holes bored 4 inches apart
in opposite directions through the shaft and bar. These joints
are satisfactory in most soils, but we have broken the socket
pins in four augers and twisted the auger bottom off in one
instance. After repairing we had no further trouble. Since
losing one twisted auger bottom under 18 feet of water we
have had the joints made stronger by screwing a nut on the
bar which extends through the socket in the arch (fig. 1, &).
We have had only one socket pin break in forty-eight augers
since using the nut and in this instance the shaft revolved free-
ly in the socket, but the nut on the underside stopped the auger
bottom from coming off. The auger struck rock and was sud-
denly stopped. The blades of the auger were bent, which shows
the strength of the joint. In the Straits Settlements in one
area we bolted iron plates to the shaft and arch to take the
strain off the socket, but at that time we did not use the extra
nut.
We are now making the joint by welding the socket, bar,
and shaft together at a cost of 8 pesos less per auger than the
cost of boring the holes and using bolts, pins, and the nut. We
have never had a welded joint break.
Turning the auger. — A simple and cheap way to turn the
auger is to hitch a rope around the shaft and use a bar of
wood or metal as a lever, but in our experience we find turn-
ing handles designed for the purpose are more satisfactory. The
Wilson wrench, one of the best on the market and commonly
used by oil drillers, is an excellent device for turning, but these
wrenches are comparatively expensive. A more satisfactory
wrench for latrine boring and used by us in the Straits Set-
tlements and the Philippine Islands costs 7.50 dollars complete
and has many advantages. This wrench is made by bolting a
handle made locally by a blacksmith to a 2^-inch Vulcan bijaw
or similar chain tong usually carried in stock by hardware
dealers. The additional handle greatly facilitates turning, and
by slipping a 4-foot length of pipe over each handle the leverage
is greatly increased. This wrench can be readjusted to any
position on the shaft without removing the chain, by a quarter
46, 4
Yeager: Bored-hole Latrines
jrn £2L_jiil
687
^33 rarnor
Fig. 2. The chain-tong turning handle, made by attaching the handle a to a Vulcan bijaw
chain tong. This is an excellent turning device and has been used in many places.
FIG. 3. Crumbie tongs, one of the most satisfactory inexpensive tongs on the market.
a, Old type ; 6, improved Crumbie tongs ; these cost a little more, but are worth it.
reverse turn and sliding the wrench up or down. Fig. 2 shows
the simple construction of this wrench.
One of the most satisfactory and inexpensive wrenches is the
Crumbie or the improved Crumbie wrench shown in fig. 3.
This wrench does not damage the shaft and costs only about
3.50 dollars United States currency. An extension handle sim-
ilar to the one shown on the Vulcan bijaw greatly improves
the use of this wrench for boring. The National Supply Com-
pany and many other dealers carry these wrenches and the Vul-
can bijaw in stock.
688
The Philippine Journal of Science
-Shaft f'pfpe
1931
Uppepp/ate
lower j/ee/
refer?/ /or? p/afe
Threaded fop
/"p/pe hand/es
Fig. 4. Turning handle designed by Doctor Hamilton. "The two steel retention plates
with the grooves on their inner surfaces serve to prevent the dogs from falling out of
place when no pipe is between them. The plates are bolted together and to the frame of
the body so that the dogs are easily removable and exchanged when worn or damaged."
Another wrench, shown in fig. 4, was designed by Dr. A. H.
Hamilton in Java. He has this wrench made locally, but the
cost, 40 pesos in the Philippines, is too high for general use.
Pipe-cross turning handles. — For economy and simplicity we
designed the turning handles shown in fig. 5, a. We first used
these handles in the Straits Settlements, and since making mi-
nor improvements we are using them on most of the augers in
the Philippines. These handles are easy to assemble and can
be made easily. A cross joint, or four- way water-pipe joint
as it is sometimes called, large enough to slide up and down the
shaft, two 4-foot pieces of pipe, and a f-inch chrome or tool-
steel pin are all the parts needed. Since breaking a few of
the light-weight crosses usually used for water pipe, we use
heavier steam-pipe crosses. A hole is drilled through the cen-
ter of the cross, through which the f-inch tool-steel pin can be
46, 4
Y eager: Bored-hole Latrines
689
o
Cross-
'5" hole
Turning
handle
Aujer arch
szz
rma N^/t»v il^j.
Fig. 5. Turning handles, a, The pipe-cross turning handle. This outfit is usually used
in the Philippine Islands for general distribution because it is the least expensive
satisfactory device we have tried. The only materials required are two pieces of pipe,
a heavy cross or four-way joint and a tool-steel pin to transfix the cross and shaft.
The cross slides easily up or down the shaft; b, a turning handle for use on a shaft
drilled with holes to engage the lugs. This wrench is more expensive than the pipe-
cross handle.
inserted. At times we ream out the cross to make it slide easily
on the shaft. The shaft is drilled at 18-inch or 2-foot inter-
vals so that the position of the cross can be changed when nec-
essary. The two 4-foot pipes are screwed into the cross on
opposite sides of the shaft and serve as levers for turning the
auger.
The cost of this cross turning handle depends upon the cost
of material and labor. In Manila the auger shaft drilled and
welded to the auger bottom and the cross turning handle, com-
plete with pin ready for use, costs 17.50 pesos delivered to our
690
The Philippine Journal of Science
1931
storeroom. Another handle is shown in fig. 5, b. This handle
hooks in a hole in one side of the shaft and a screw enters a
hole on the opposite side. This handle costs double the price
of the cross handle.
The coupled shaft. — Where the transportation of the 20-foot
shaft is difficult, a coupled shaft made of two 11- or 12-
foot pipes can be used. A strong joint that will not wabble
can be made by riveting a round solid iron bar 12 inches long
into the end of one section of the shaft, allowing 6 inches of
the bar to extend out of the end as shown in fig. 6, a. The
protruding bar and the end of the other section of the shaft
can be drilled in opposite directions for the insertion of two
f-inch bolts, which can be removed when transporting the au-
ger. If only one cross turning handle is used the upper sec-
tion of the shaft will have to be added every time a latrine
reaches a depth of about 11 feet. This takes only a minute,
but if a ^number of latrines are to be bored in one area it saves
time to have two crosses, one below the bolts and one above
the bolts on the shaft. The 4-foot handles can be unscrewed
from the lower cross and screwed into the upper cross by hand
when necessary to shift to the upper section of the shaft. Each
cross allows the handles to be shifted 11 feet. Another
method is to bolt six crosses permanently to the shaft at 3-foot
intervals, then the handles can be screwed in each cross as the
boring becomes deeper, without shifting the cross. Our work-
men prefer shifting the cross, and six crosses increase the cost ;
therefore, we use only one or two crosses.
£
§ piVefs
^At/gep shaft
Lowep sect/on
ysBqtt hotes^
) I
Mi'/d steet hap
a
*sf
Augep shaft*
Uppep section
1
-w-
-**-
~
S3 £=2
-*PQ-
ittr
ntir
Fig. 6. Coupling and shaft, a, A solid coupling for shafts made in sections ; two bolts
are removed to take the shaft apart ; 6, the bolt shaft, one of the most useful shafts
we have used. This shaft stands more rough use than any other shaft tried. Permanent
bolts or rivets transfix the shaft at 3-foot intervals. The turning handles never damage
this shaft. The bolt heads act as lugs for the handles to push against.
46,4
Yeager: Bored-hole Latrines
691
The bolt shaft and turning handles. — This method of turning
is simple to use, fool proof, and easy to make. The bolt shaft
shown in fig. 6, &, is made by inserting a series of pins or
bolts through the shaft at 2-foot intervals, allowing the head of
the bolt and nut to serve as lugs. If the bolts are too long
cut them off flush with the nut. The nuts on the bolts are
screwed against the shaft tightly and are never removed. The
heads of the bolts and the nuts serve as lugs for the turning
handle to push against to prevent the handle slipping around the
shaft, and does away with the use of expensive chain tongs
and wrenches, which damage the shaft to some extent. The
turning handle is made from an iron bar, which is shaped so
that it will hook on the nut on one side of the shaft and push
on the bolt head on the opposite side and can be quickly attached
for turning. Two different turning handles are shown in fig.
7, a and b. This device is a great time saver as there are no
clamps, chains, screws, pins, or ropes to adjust. In one in-
stance the shaft of an auger equipped with the cross turning
Fig. 7. Turning handles, a, One type of locally made turning handle for the bolt shaft,
hammered out by a blacksmith. These can also be made of cast iron ; b, another type of
locally made handle for the bolt shaft. The head of a bolt on the shaft enters the
socket r, and the nut on the opposite end fits into notch n; c, a turning handle that
can be made locally or purchased ready-made from dealers.
692
The Philippine Journal of Science
1931
Section of
-frpipe
ruminq handle
■ W, Auger
shaft
WA
3fmf 4%h
Turning handle
Fig. 8. A more elaborate turning handle for use on the bolt shaft. This is an excellent
handle but costs 20 pesos to make locally. In large quantities the cost would be leBS.
46, 4
Y eager: Bored-hole Latrines
693
handle was damaged so that the cross would not slide up the
shaft, but the permanently bolted shaft stands a great deal of
damage before it is put out of use. If the projecting bolts are
objectionable, one bolt can be used and shifted from hole to hole
when necessary to change the position of the handle. The se-
ries of permanent bolts are suggested in order to save time.
The handle shown in fig. 7, c, is useful if rivets are used instead
of bolts to prevent slipping. A more elaborate wrench, shown
in fig. 8, has no damaging teeth and can be made locally. There
are many excellent ready-made wrenches that can be purchased
from dealers, but these are more expensive.
Fig. 9. The A frame now used instead of a tripod for hoisting augers. The guy ropes are
usually tied to a house, tree, or fence post. Bamboo is usually used because it is
much cheaper in the Philippines than iron pipe.
694
The Philippine Journal of Science
1931
Fig. 10. In some places augers are lifted
by direct pull, but the job is too heavy
in most areas. The rope is attached to
the arch or low down on the shaft.
HOISTING DEVICES
Tripods. — A tripod 25 feet in
length made of bamboo or other
wood or 2-inch water pipe is ve-
ry useful for hoisting the auger
by means of a pulley and rope.
Inverted V- or A-shaped
frames. — For the past two years
we have discontinued the use of
tripods and now use only two
poles or pipes supported by two
guy ropes (fig. 9). The A
frame is easier to transport and
erect than a tripod and can be
put up in places where there is
not enough space for a tripod.
The guy ropes can be attached
to a house, trees, posts, or stakes
driven into the earth. In some
localities we get the bamboo
poles for nothing, and in other
places the bamboo costs 3 pe-
sos or more. Bamboo breaks
after using it a few times, and
in places where it cannot be ob-
tained or is expensive we make
an inverted V frame from two
li-or 2-inch pipes fastened at
the top with an iron bolt. The
feet of this A frame are two T
joints screwed on the ends of
the legs. The T joints prevent
the ends of the pipe from sink-
ing into the earth. A stick or
bar can be inserted into the T
to prevent sinking in very soft
soil.
Pulleys. — We have used a num-
ber of 12-inch well pulleys or gin
46,4 Y eager: Bored-hole Latrines 695
blocks for hoisting, but now use 4- to 7-inch compound pulleys
of wood or metal. We usually use a double 6-inch pulley at the
top of the frame and a single 4- or 6-inch pulley to attach to
the auger. Compound pulleys are so well known that a descrip-
tion is not needed in this article.
If the soil and auger is not too heavy a tripod or hoisting
frame is not necessary and the auger can be lifted by a direct
pull on the rope attached to the arch of the auger as shown
in fig. 10. It is difficult to start lifting an auger full of earth,
but when once loosened it is easy to pull to the surface. The
initial lift can be done by direct pull on the turning levers
or by placing a plank or two across the mouth of the hole and
applying leverage. Planks can be placed across the mouth of
the hole and be notched so that they will support the shaft
while turning.
The direct-pull method is too heavy in most areas in the Phil-
ippines, where we have even stopped using single pulleys and
use compound pulleys. Dr. John L. Hydrick states that the di-
rect-pull* method is used successfully in Java and there is no
tripod to transport.
Shaft support. — Latrines can be bored with only an auger,
pulleys, and a tripod, but there is a great saving of energy if
some kind of brace is used for the shaft of the auger. The most
satisfactory cheap support we have used is a pair of doors,
which cost 3 dollars United States currency, including material
and labor. The doors are put in place as soon as the latrine is
bored to a depth of about 2 feet. The doors are closed over
the hole when the auger is lowered, and opened when the auger
is raised. The doors are hinged to a wooden frame about 36
inches square, and the shaft of the auger turns in a hole cut
in the closing edges of the doors (fig. 11).
Another type of brace that is cheap and satisfactory is shown
in fig. 12, a. This brace is fastened to the frame about 5|
feet above the surface of the earth. A tipping auger can be
lifted and emptied into a receptacle over the latrine, or a knock
on the clamp of the brace will release the shaft, allowing the
auger to be swung away from the hole to be emptied.
An excellent brace designed by Doctor Hamilton in Java is
shown in fig. 13. This brace should hold the shaft steady, but
the cost of making is too great for general distribution in the
Philippines.
696
The Philippine Journal of Science
36"
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Fig. 11. Trap doors costing 6 pesos greatly facilitate boring. The auger shaft is sup-
ported by the doors when closed. Stakes can be driven into the earth at the notched
corners to prevent movement of the platform while turning. One of these platform
braces is now included as standard equipment with every auger used in the Philippines.
A quadruple brace sold by the Alsdorf Corporation is shown
in fig. 14.
Accessories. — A bamboo pole or tamping rod is used in some
kinds of soil, especially certain kinds of sands, to tamp the earth
46,4
Yeager: Bored-hole Latrines
697
Fig. 12. This tripod is more difficult to transport than the A frames, but if properly
made greatly speeds up boring: when several holes are to be bored within a small area.
a, Shaft support, A knock on the extension lever c releases the auger so that it can
be swung away to be emptied; b, an iron hook that is a great time and energy saver.
These hooks hold the auger away from the latrine while being emptied. The hooks are
also used on A frames.
more solidly in the auger to prevent it from falling out when
lifted. A little clay can be thrown down the hole in some places
to make the earth more cohesive. Clay is sometimes used in
26420!
698
The Philippine Journal of Science
1931
well drilling in very soft earth to prevent temporarily the sides
from caving.
A long piece of 1- or lj-inch iron pipe with a tool-steel chisel
riveted or welded into one end is useful for breaking hard strata
or straightening the sides of the hole. Ordinary straight chi-
sels, the T, or cross chisels, can be used.
An inexpensive swivel to prevent the ropes twisting can be
used on top of the auger shaft,
but this requires a high A
frame. A swivel that will
screw on the end of the shaft
is made from an ordinary cap
and iron ring as shown in fig.
15. Even a 15-foot A frame
f\
Fig. 13. An excellent shaft brace designed
by Doctor Hamilton. The cost of pro-
duction is a disadvantage of this device.
(Drawing from Doctor Hamilton's
sketch. )
Fig. 14. A quadruple expansion brace
sold by the Alsdorf Corporation. The
stock size is made to fit the Standard
earth auger.
46,4
Y eager: Bored-hole Latrines
699
can be used, if a removable turn-
ing handle is used and no swivel,
by hooking the lower pulley of
the tackle to a movable iron ring
on the shaft. We usually use a
25-foot A frame for greater con-
venience, and on augers equipped
with the cross turning handles,
which are not removed when
hoisting, we provide an iron
ring that can be shifted and
costs 40 cents at a ship chand-
lery or hardware store. The
bolt of this ring is slipped into
one of the easily reached holes
of the shaft, and the tackle is
hooked on each time before lift-
ing. It is unhooked when turn-
ing the auger to prevent twist-
ing. Devices for hooking the
tackle to the shaft are shown in
fig. 16.
GE
x:
^§^ ii'tnQs
R/ng
a
Nut
*-Cap
Fig. 15. A locally made swivel for use on
the top of the shaft. We have used
these swivels but prefer unhooking the
rope from the shaft while turning the
auger.
i#m
6
EiSii
Fig. 16. Devices which can be attached to the auger shaft and adjusted to any position
so that a hook on the hoisting rope can be quickly hooked on instead of tying and
undoing knots; a and b are used on shafts with holes drilled at intervals such as
used with the pipe-cross handles ; c and d are used on the plain or bolt shafts.
700 The Philippine Journal of Science issi
a
b
Fig. 17. Reamer, stone hooks, and grapple, o, An undercutting reamer which is useful
in cutting away the sides of latrines below linings ; fortunately it is rarely necessary
to use one of these; 6 and c, stone hooks which are useful in removing small bowlders;
d, a grapple (redrawn from picture from R. R. Howell & Co.).
We find an iron hook (fig. 12,
6) attached to a leg of the A
frame very useful to hold the
auger away from the latrine
when emptying. This saves
energy because a man does not
have to hold the auger for an-
other man to empty.
The stone hooks and grapple
shown in fig. 17, b, c, and d, are
useful for hooking and pulling
large stones out of the latrine.
The undercutter or reamer
(fig. 17, a) is useful for cutting
away the sides of the latrines
below linings to facilitate sink-
ing cylinders.
A cheap boring equipment for
regularly hired squads. — The
apparatus shown in fig. 12, while
probably too expensive for gen-
eral distribution, greatly facili-
tates rapid boring for the use
Fig. 18. Augers suitable for soft mud and
silting sand, a, Type made by many
manufacturers of drilling equipment; b,
a heavy dumping auger such as used
with machine-driven outfits; this auger
is made by the Gus Pech Co.; c, the
Lang auger with sand-boring screw;
this is a good hand auger for boring in
sand. See fig. 31, Howell drop-bottom
auger.
46,4
Y eager: Bored-hole Latrines
701
Pig. 19. Sand pumps and bailers which can be purchased from dealers. These pictures
are from the Keystone Drilling Co., Beaver Falls, Pa.
of a squad of regularly hired men. Four men can work faster
with this outfit than any other equipment we know of at the
price. It will be noted that two legs of the tripod are close
together to allow use in limited space, and to furnish extra sup-
port where there is the greatest strain. The tripod is made
702
The Philippine Journal of Science
1931
door Aflm
View
Boons
QqqP locally of 2-inch water pipe. The cost of
the equipment in Manila is 54 dollars for
the tripod complete with winding drum
and auger brace, not including the auger.
This apparatus is more difficult to trans-
port than the A-frame outfits.
Boring soft sand and mud. — These ma-
terials can be forced out with a pump, but
it is not as satisfactory in routine work
7hn V/PW as an auger esPed&Uy designed to do the
* work. A number of excellent sand bailers
and augers sold by dealers are shown in
figs. 18 and 19. Some of these work by
a pounding motion, percussion, or spud-
ding, and others work by rotation. Those
shown in fig. 19 are not very useful in
latrine installation. An excellent cheap
rotating sand auger can be made by add-
ing valves, or what might be called trap
doors, to the ordinary Iwan post-hole
auger. These doors are shown in fig. 20.
Soft sand falls out of the sides of this
auger as. sent out by the manufacturer,
but a piece of sheet metal hinged to the
noncutting sides of this auger so that it
will open about 2 inches, allowing sand to
enter but not to fall out, serves the pur-
pose in some places. Exceptionally soft
sand will fall out of the bottom even when
fitted with side valves, and in these places
two additional doors should be fastened
to the blunt edges of the blades crossing
the bottom of the auger. The auger altered in this fashion
does the work but is not as satisfactory as augers especially
designed for silting soil. In order to empty this auger the shaft
must be lowered almost horizontally, or a tipping hinge on the
shaft can be made like those shown in figs. 21 to 23, which
are made locally for use in the Philippines. These hinges al-
ways stop on a dead center, allowing the locking pin to be in-
serted quickly. A tap with a hammer or block of wood knocks
the pin out when necessary. To facilitate transportation the
Fig. 20. The Iwan post-
hole auger fitted with
valves for use in silting
sand.
46,4
Yeager: Bored-hole Latrines
703
Fio. 21. A hinge to facilitate turning an auger over so that it can be dumped.
704
The Philippine Journal of Science
1931
Auger
shaft
Auger
arch
Fig. 22. A hinge to facilitate turning an auger over so that it can be dumped.
46,4
Yeager: Bored-hole Latrines
705
Fig. 23. A hinge for the same purpose as those shown in figs. 21 and 22.
auger bottom can be removed
from the shaft by taking out the
main bolt. Another type of
hinge, used on an auger de-
signed by Mr. A. L. Savignac for
the United Fruit Company, is
shown in fig. 24.
fig. 24. An auger designed by a. l. sa- An excellent auger can be
vignac for the United Fruit Co. This ma(Je locally aS Shown by fig. 25.
auger works in soft mud. Note the __ - _ _ „ . . , ,
hinge for dumping. The blades of this auger can be
706
The Philippine Journal of Science
1931
ii
I, 35 cm d/a. rl
Section A -A
Fig. 26.
A valve auger designed by Doctor Hamilton
in Java.
Fig. 25. A clay and sand auger
that can be made locally.
Augers of similar design are
sold by many manufacturers
without the flap valve a. R.
R. Howell & Co., Minneapolis,
manufacture these augers. A
hinge on the shaft is not
needed because the bottom of
the auger swings back on a
hinge to empty the contents.
opened quickly by hit-
ting the catch, allow-
ing the bottom with
blades to swing on a
hinge out of the way.
Doctor Hamilton designed the auger shown in fig. 26. We
have not given this auger a trial and are not prepared to re-
port upon its efficiency.
The engineers of the Sarawak Oil Fields designed the auger
shown in fig. 27, but this auger costs 65 dollars to make locally
without shaft or other accessories. It requires six men to handle
it and no doubt is useful if properly handled.
OTHER CLAY AND SAND AUGERS
There are many kinds of these augers on the market. A style
frequently used a number of years ago and still used in many
places is the disc auger (fig. 28). We have tried a number of
these augers but find them not as satisfactory as the other au-
gers described.
46,4
Yeager: Bored-hole Latrines
707
m
<o
<o
10" >
pip*
Sect/oh
Scale I"=IFoot
Scale 3"= I Foot
Plan
Fig. 27. A short chisel-bottom bailer designed by the engineering department, Sarawak
Oil Field, Ltd., Miri, Sarawak. This drill should be a good one, but is expensive to
make and requires six men to handle effectively.
The Standard earth anger (fig. 29). — In some places these
augersTwhich are equipped with extension bits, have been used.
They are fine augers for making holes of small diameter and have
the advantage of being made so that the blades open to f acili-
708
The Philippine Journal of Science
1931
tate dumping. These augers give excellent service in certain
kinds of work, but after a thorough trial in the Straits Settle-
ments in making holes large
enough for bored-hole latrines
we decided in favor of other
augers. The Iwan post-hole
augers for instance cut 14- or
16-inch holes more rapidly in the
variety of soils we encountered.
The Lang borer (fig. SO). —
This auger is used in a number
of countries, but in early trials
it did not meet with much suc-
cess in the soil in the Straits
Settlements. Holes were cut
with the Iwan auger in one-third the time required in the same
soil with the Lang borer. The handles and extension rods of
the Lang borer are likely to bend, and there is a great waste of
time using the coupled joints recommended by the manufac-
turer and in using the lifting bars sent with the auger. The
suction created in some soils when lifting made this job diffi-
cult compared to other augers. An advantage of the Lang au-
ger as equipped by the manufacturer is that it will work suc-
cessfully in soft sand, and the Iwan post-hole auger will not
work in very soft sand without modification. The Lang 14-
inch auger with deep boring attachment costs 7 pounds 8 shil-
Fia 28. Disc .augers. These are probably
the cheapest angers made, but are not as
good as other augers mentioned.
MOVABLE BLADE WHICH
OPENS TO DUMP EARTH
CLOSES AGAIN AND
LOCKS Of ITS OWN
WEIGHT
TRIGGER WHICH RE-
LEASES MOVABLE
BLADE WHEN SLID-
fNG HAND LOCK IS
STANDARD SLIDING
HAND LOCK. WHEN
THIS IS RAISED. TRIG-
GERISRELEASED AND
MOVABLE BLADE
OPENS WIDE TO
OUMP
HOLES IN STANDARD
EXPANSION BLADES.
_ THESE PERMIT AD-
JU8TMENT8 FOR
BORING DIFFERENT
SIZE HOLES.
SINGLE THUMBSCREW
FOR ADJUSTING
STANDARD EXPANSION
BLADE TO CUT HOLES
UP TO 16 IN IN OIAM.
ETER
NOTE THE CURVE OF THE BLADES StE
- HOW THEY BORE THE FULL WIDTH OF THE
HOLE AT THEIR POINTS
Fig. 29. The Standard auger, sold by the A. J. Alsdorf Corporation.
46,4
Y eager: Bored-hole Latrines
709
lings 9 pence, and with five extension rods, two levers, one
spiral joint, and one steel chisel, costs 19 pounds 6 shillings 3
pence. A report recently re-
ceived states that the Lang
auger is now made with rods
and handles heavy enough to
stand the strain of deep latrine
boring. If the manufacturer of
this auger used a larger one-
piece shaft and longer turning
bands the cost could be reduced
as well as the time required for
boring. The coupled shaft is
unnecessary for latrine boring
in most places. The shaft as
supplied can be taken apart for
shipment, but this is not a great
advantage, because an auger is
usually used in one area for a
long time and carried f romj house
to house completely assembled.
Howell's augers. — A variety of augers manufactured by R. R.
Howell & Co., Minneapolis, Minnesota, and used by drive-well
men are shown in fig. 31, a to /. These augers are too long
and heavy for a squad of four men to handle. The spiral auger
for loose sandy soil shown in the same illustration is carried in
stock by the manufacturer in sizes up to 16 inches in diameter.
The worm of this auger is 4 feet long. We have not tried this
auger, but it would probably be more satisfactory for latrine
work if made only half this length unless several men or a
power-driven machine is used. The 12-, 14-, and 16-inch dia-
meter augers cost 40, 45, and 50 dollars, respectively. These
augers should be worked through a casing, if the sand does not
pack tightly enough to keep it from running out.
The drop-bottom auger shown in fig. 31, g, has been used
for years by well drillers. It is an excellent sand and clay
auger but heavy, and the 16-inch size costs about 50 dollars.
The spudding jet auger shown in fig. 31, h, is for rock drill-
ing but is slow and requires a heavy rig. It is used in well
drilling but dynamite is faster and cheaper in latrine instal-
lation.
Fig. 30. The Lang-London, Ltd., auger.
This auger is used in many places in
clay and especially in soft sand. When
used in sand a special screw, shown in
fig. 18, c, is attached.
710
The Philippine Journal of Science
1931
Fig. 81. Various earth augers manufactured by R. R. Howell & Co. a, For clay and
hard pan ; b, for boring and removing core ; e and d, for general boring ; e, for loosening
and removing stones; /, for loose sand soil; g, a drop-bottom, fast-cutting auger
especially useful with power-driven machines; h, a spudding, jetting drill used in rock
drilling. Blasting is much more rapid for latrine installation in rock.
GEARED AUGERS FOR MAN POWER
Geared apparatus that can be turned by hand is made by a
number of manufacturers, but the speed gained in drilling does
not justify the expenditure for latrine boring and the apparatus
is more difficult to transport and set up than the apparatus
described in the first part of this article. The geared drills for
making the 1-inch blast hole are worth the money. The geared
hand augers made by Ingersoll-Rand for cutting 16-inch holes
cost about 1,750 dollars United States currency.
ANIMAL-DRIVEN AUGERS
If horses, bulls, or other animals are available heavy rotary
or studding drills can be used. These outfits are shown in fig.
32. In most villages where latrines are to be installed there
is not enough working room to rig up apparatus of this kind,
and by the time the outfit is set up a hole made by laborers
would be well under way. Animal-driven boring apparatus is
46,4
Yeager: Bored-hole Latrines
711
Fig. 32. Animal-driven boring apparatus. Used for many years in well drilling, but not
suitable for rapid latrine boring.
valuable for deep-well drilling, but is not practical for 20-foot
latrines; therefore, the heavy equipment used for this purpose
will not be described in this article. Heavy augers, the bits of
712
The Philippine Journal of Science
1931
which cost about 60 dollars each, have been designed for eight
men or animals and do not speed up latrine boring enough to
justify using them.
POWER-DRIVEN MACHINES
A power-driven auger to be of practical value must be small
and easily transported. There is no doubt about the efficiency
of these machines, and there are plenty of statistics to show
that power-driven holes can be made for less money per hole
than by man power in places where a large number of holes
are to be bored.
BUDDA-HUBRON EARTH DRILL
One of the most compact, easily transported, rapid boring
machines is the Budda-Hubron auger shown in fig. 33. For la-
trine boring this is the handiest and one of the most efficient
machines on the market. It will bore in nearly all soil forma-
tions including shale, frozen ground, and hardpan. The appa-
ratus usually sold for post holes bores to a depth of 10 feet.
Fig. 33. The Budda Hubron machine drill. This is an efficient rotary drill, amd takes
less operating space than any machine we have heard of. It can be used for boring
in clay, sand, hard pan, shale, and frozen ground. The stock machine bores holes
10 feet in depth, but the manufacturers will equip it for boring 20-foot holes. It costs
about 2,400 dollars United States currency.
46,4
Yeager: Bored-hole Latrines
713
With special equipment this machine will bore a hole 20 feet
deep. The machine can be equipped with small drills for making
blast holes or holes up to a diameter of 24 inches. The manu-
facturer has certified records showing that this machine sunk
fifty-four 7-foot by 22-inch holes in clay and gravel in nine
hours at a cost of 0.454 dollar per hole. Hand methods cost
2.70 dollars per hole. Many other records are available.
THE GUS PECH MACHINE
The apparatus shown in fig. 34, manufactured by the Gus
Pech Foundry and Manufacturing Company, requires an oper-
ation space of 10 by 16 feet, which rules it out for latrine in-
stallation in most places. The machine will bore from seventy
to eighty post holes a day in some soils and will work in any
kind of soil free from rocks. The machine is powerful enough
to handle a 24-inch auger, and a reamer to cut 36-inch holes.
There are some disadvantages in constructing these large diam-
eter latrines, but if boring in an area where numerous large
bowlders are encountered it is easier to remove these obstruc-
tions than to try to bore through them. A 16-inch drop bot-
Fia. 84. The Gus Pech power-driven machine is a rapid borer, but requires a space 10
by 16 feet for efficient operation. It costs less than 1,000 dollars equipped for latrine
boring.
264209 12
714 The Philippine Journal of Science 1931
torn, double-bit auger sells at 45 dollars. The 24-inch auger
with reamer costs 60 dollars. Equipped to bore 16-inch holes
20 feet deep, a 6-horse-power gasoline engine, and a number of
accessories, the machine sells for 842.50 dollars. There is an
additional charge of 145 dollars for exportation boxing.
THE KEYSTONE, MONITOR, STAR, AND OTHER MACHINES
There are many machines, including those mentioned, on the
market that I assume could be adapted to latrine boring, but
I do not know of any better suited to the purpose than those
described. All of the manufacturers making these machines
deal in a large variety of augers and accessories. A disad-
vantage of all these machines is the large space required for
operation. The Budda-Hubron requires less space than any
power-driven machine that has come to my attention.
BORED-HOLE LATRINES IN ROCK
Expensive power equipment will cut through rock without
difficulty, but to suit Philippine conditions it was necessary
to develop an inexpensive method of latrine installation, be-
cause there are many towns near Manila and in other parts
of the Islands built on strata of tuff. In an article submitted to
the Rockefeller Foundation the formation, correctly named tuff,
was referred to as adobe rock, as locally termed. Tuff in the
Philippines is found in several degrees of hardness.
The softest tuff can be cut with a pick, but the hardest grades
break into sharp-edged irregular pieces when blasted with dy-
namite, but is not as hard as granite or the solid rock forma-
tions met with in some places. The so-called adobe rock in the
Philippines does not melt away in the rain as adobe does in
many countries. This tuff stands weathering for centuries as
seen in some of the old unprotected walls and churches. The
harder formations of tuff offer more resistance to boring than
the laterite frequently encountered in Malaya or the hardpan
that is found in other countries. Tuff is a deposit of lava and
volcanic ash that by pressure and other causes has hardened and
forms extensive strata varying from a few inches to 30 feet
or more in thickness.
Many towns are built on outcrops of this rock, where the
pail system and other methods of disposal of dejecta were too
expensive; therefore, such places had no latrines.
46,4
Yeager: Bored-hole Latrines
715
The problem of making holes economically in these areas
seemed impracticable until the method of making tunnels for
railways and water-ways led to the idea of making miniature
tunnels vertically instead of horizontally. The method is the
same in both instances. Dynamite does the work faster and
cheaper than any auger made. In our first attempt we blew
the surface of the earth to pieces several yards around the mouth
of the hole, as shown in Plate 7, fig. 1, but with a little modifi-
cation in the method, cylindrical holes with clean-cut mouths
and straight sides can be made to any depth desired. We
have installed hundreds of these latrines within a few feet of
the houses. Some of these holes have been blasted within 2
feet of the walls of the houses and others have been put down
under the floors. Any person who understands the use of dy-
namite can install these latrines in crowded communities without
any danger to the inhabitants or houses.
The method of blasting varies according to the kind of rock
encountered. Some of the tools employed are shown in fig. 35.
Fig. 35, aS and a4, shows two views of a drill for making blast
holes in hard rock. This drill is not needed in adobe forma-
tions.
£> «20
Fig. 35. Tools used for blasting latrines in rock, a, Bar used in drilling blast holes
in tuff, adobe rock, and other hard formations ; a S, drill used with hammer for making
blast holes in hard rock ; b, crow bar used for starting latrine or straightening side ;
c, long bar used occasionally in deep latrines ; d, bamboo bucket for removing water
from blast holes ; c, bamboo brush for cleaning mud out of blast holes.
716 The Philippine Journal of Science ^zi
Making holes in tuff. — If the stratum of the tuff lies 2 feet
or more below the surface of the earth, the ordinary post-hole
auger or shovels are used to cut the hole through the sandy-
clay earth down to the level of the hard layer. When this
layer is reached, three to five small holes (fig. 36) about 1 inch
in diameter and 2| feet deep are drilled with the steel bar
shown in fig. 35, a. Each of these holes can be drilled in ten
to fifteen minutes by using the bar with a ramming and twisting
motion after pouring a little water into the hole. A hammer is
not used except in very hard rock. The water and mud that
accumulate in the holes are removed with the bamboo bucket
shown in fig. 35, d, and the holes are cleaned out with the bamboo
brush, e. The bucket, d, can easily be fashioned from bamboo
with a pocket knife, and the brush is made by pounding the
end of a bamboo pole. The position of the holes, the direction
of drilling, and the charge of dynamite to be used in each,
depend upon the work to be done. One blast hole in the center
of a large 16-inch hole with a large charge of dynamite is not
satisfactory if the large hole is shallow, because the blast will
destroy the surface. Better results are obtained in making 16-
inch cylindrical holes if three to five small holes are drilled in
the position indicated in fig. 36, a. A charge of one stick of
|-inch 40 per cent dynamite in each small hole is sufficient. The
fuses are all ignited as rapidly as possible. The explosions
follow each other in rapid succession.
When the latrine is 4 to 6 feet deep, the dynamite will break
up the adobe or pulverize it to an additional depth of 3 to 5
feet below the bottom of the blast holes. If the adobe pulve-
rizes an auger is used to remove it, but if it breaks into pieces
a bucket and rope is used. A man can be let down into the
hole to gather up the larger pieces of rock. We make latrines
at least 16 inches in diameter if men must go down. At this
stage after the adobe is removed it will be found that the latrine
is from 9 to 11 feet deep. Another series of small holes (fig.
36, c,) with another charge will be enough to make a latrine
pit about 18 feet deep. We frequently drill three or four blast
holes at the first level, four at the second level b, and five for
the last detonation at level c, placing one or two sticks of dyna-
mite in each hole.
If the adobe is an outcrop and not covered with soft earth, a
hole about 16 inches in diameter and 18 to 24 inches deep should
be cut into it with a chisel-shaped bar (fig. 35, 6). If the rock
is very hard the work may be expedited by a few light charges
46,4
Yeager: Bored-hole Latrines
717
wrrmwnzMx&®L
. ...!.#?■.•. . ...... ... .
1 ' u • • " . - - • •■.•..•„;■:
Fig. 36. Blasting in rock, a to e, Positions of blast holes. In hard rock five blast holes
are drilled at a and 6.
of dynamite ; but this is not often necessary even in the hardest
tuff. When the large hole is 1£ to 2 feet deep, the work of
charging the small holes with one stick of dynamite each can
proceed without damage to the surface. Sometimes the dyna-
718 The Philippine Journal of Science 1931
mite will blast out a wide hole beneath the mouth of the small
hole, and leave a shelf in the latrine. This shelf can usually
be broken through with the long chisel (fig. 35, c) or, if neces-
sary, a small charge of dynamite may be used.
Dynamite does not blow long fissures in comparatively soft
tuff, but usually pulverizes it so that it can easily be removed
with an auger. Occasionally, broken pieces several inches long
and of irregular shape are blasted loose. The dynamite can be
set off with the usual fuse and cap, but if much work is to be
done it would probably be better, in the long run, to use an
electric machine for this purpose.
In most places we have made straight cylindrical holes 18 to
20 feet deep. In other places we have first made holes 15
inches in diameter and 6 or 8 feet deep, and then, by drilling
two small holes at an angle (fig. 36, d and e), have made the
large hole 3 or 4 feet in diameter down to a depth of about 10
feet. At one school we constructed a series of five holes 3 feet
from center to center, and then blew out the partitions at the
bottom, thus connecting the holes by an opening large enough
to permit a man to walk from one hole to the other. In other
places we have connected and installed pipes in the partitions so
that the holes acted as a septic tank. There was sufficient ab-
sorption in the holes in soft adobe that a pipe to carry off effluent
was not needed.
Holes in the hardest tuff will give better service if the area
around the bottom is split into fissures to allow greater absorp-
tion. This is done by drilling one blast hole in the center at
the bottom of the latrine and setting off a charge of several
sticks of dynamite. This will blast numerous fissures several
feet long in all directions. If the rock is not then sufficient-
ly absorbent, two latrines should be made so that the dejecta
can be allowed to age in one hole while the other latrine is being
used. It is easier to pump out a full latrine than to bore a
new one in rock. A machine and hand-driven pump are used
in the Philippines for pumping out latrines.
We have used from two to twelve sticks of dynamite in making
holes in various kinds of tuff. In a large school latrine an aver-
age of seven sticks per hole was used. With the tools men-
tioned, two men can install a latrine in tuff in a day or less.
Dynamite costs 24 pesos, or 12 dollars United States currency,
for a case of two hundred sticks, delivered in Manila. In hard
tuff the dynamite for one latrine costs from 42 to 72 cents. The
46,4
Y eager: Bored-hole Latrines
719
cost of installation, while higher than boring in soft earth, is
cheaper in most places than any other method of latrine con-
struction in hard formations. Dynamite and labor cost about
4.50 pesos per hole if only twelve men are hired to lower the
overhead cost per hole. The foreman's wages of 3 pesos per
day are included in the cost.
The cost can be considerably lowered by spending a little
more money for tools. We had a special blast-hole drill made
by the Howells Mining Drill Company that speeds up the work.
It not only bores the blast hole but automatically cleans the
hole while boring. This auger, shown in fig. 37, costs 18 dollars
delivered, including an extra twist drill. The drill will cut adobe
rock, shale, laterite, hardpan, and other hard formations. It
will not cut the hardest grades of tuff. A more-rapid autom-
atic feed, geared drill that costs 140 dollars is shown in fig. 38.
Air and electric drills are much faster and will cut hard rock
but are expensive unless there is a great deal of work to be done.
L_o_j£ — » °i
30'
o d
3C
liT-ol
10"-
Z3=
3 C
*/■ o »
Fig. 37. A hand drill especially designed for making blast holes in tuff, adobe rock,
shale, or other hard formations. It will not cut hard rock. This drill cuts the holes
rapidly and cleans them out at the same time ; it is constantly used now in place of
the bar a, fig. 35 except in the hardest rock where the drill a3 is used. Manufactured
by the Howells Mining Drill Co., Plymouth, Pa. Cost about 18 dollars.
720 The Philippine Journal of Science 1931
Dynamite exerts a more powerful explosive force in hard
formations than in soft material, and the hardest rock cannot
resist a charge of dynamite. While this method has not been
used for our latrine installation in the hardest kinds of rock,
it is believed that the method will work in any rock formation,
and in any formation is much more rapid than the spudding
rock drill shown in fig. 31, h, or any other drill designed for hard
formations.
ESOSfrCSXtt
Fig. 38. A geared drill for boring blast holes rapidly in very hard formation except hard
rock. Manufactured by Howells Mining Drill Co., Plymouth, Pa. Cost, 140 dollars.
LATRINE CONSTRUCTION
The method used in making holes in rock might have been in-
cluded in section under Construction but a description of the
tools used should be included under equipment, so the method
of using the tools was included with the description of the tools
in order to describe completely the method of latrine installa-
tion in rock without referring to other parts of the article.
Boring latrines in ordinary soils. — The method of using the
hand auger needs no detailed description. The auger is turned
until full and then pulled up and emptied. After a few trials
the number of turns necessary to fill the auger can be deter-
mined. The number of turns varies in different soils. The
Iwan post-hole auger takes about 6 inches of soil every time it
46,4
Yeager: Bored-hole Latrines
721
is filled. This observation is useful when boring under water
where the auger cannot be seen. A mark on the shaft can be
noted and when it reaches a level of 6 inches below the start-
ing position it indicates that the auger is full and ready to be
hoisted and emptied. Soft soil can at times be kept from falling
V\
A\\
cf*#
\v\
\v\
4
\\\\
can
1 "llir^
V
^ \/fr
5I>
"1
Fig. 39. Derrick manufactured by Werf Conrad, Haarlem, Holland. For rotary, percus-
sion, or free-ball system of boring. This apparatus will handle heavy drills. It is too
heavy for general bored-latrine work.
722 The Philippine Journal of Science 1931
out of the auger by a smooth steady pull instead of a jerky
motion when lifting. Some soils will not fall out of the auger
until after it is pulled out of the water. At times it is best to
continue a steady pull all the way up, and in other instances it
is advisable to stop lifting as soon as the auger is pulled above
the surface of the water to let the excess water drain out about
a half minute, allowing the soil to pack itself and then continue
hoisting.
Soils of average consistency pack so tightly in the auger that
they must be removed with a small scoop or sharpened paddle
made of metal or wood. Boring is easier in very dry soils if
water is poured into the hole.
Bored latrines can be installed in places where water is not
encountered, but the disintegration of the dejecta appears to be
not as rapid or complete as in latrines with a meter or two of
water in them. The dry latrines evidently do not last as long
as those containing water, but with proper use a latrine not
containing water should not be filled by an average family in less
than four years. We had a complaint in which two latrines were
reported filled within seven months, but investigation showed
that nearly one hundred persons were using these two latrines.
Ample provision should always be made to install a reasonable
number of latrines for the convenience of the persons who will
use them. There are records of other latrines lasting three
years and not yet half full. In these instances from eight to
twenty persons used the latrines daily.
A squad of four men can easily bore an average of three lat-
rines a day in sandy clay. This includes setting up the appar-
atus and time lost in transportation, and covers work by the
month, and not a spurt of energy for only a few days.
Caving soil — Lining or reenforcement to prevent caving are
important features of construction. There is one area in the
Philippine Islands where the people went ahead with bored-hole
latrine installation without preliminary consideration of the soil
encountered, and no trial latrines were bored before general in-
stallation started. Evidently in this area the soil was of a con-
sistency that the walls of the latrines did not cave in when
bored, but when the heavy rains came most of the latrines filled
up with caved-in soil. This was damaging to our propaganda
because a number of persons were convinced that bored-hole
latrines are not suitable for the Philippine Islands.
46,4
Y eager: Bored-hole Latrines
723
Fig. 40. Hand-power spudding and hoisting windlass manufactured by Werf Conrad, Haar-
lem, Holland. Believed to be too expensive (about 200 dollars) and too bulky for
general distribution for latrine boring.
In many places the earth caves in only at the mouth of the
latrine. In these areas wooden cement kegs which can usually
be obtained at no expense are used. One cement keg pushed into
724
The Philippine Journal of Science
1931
nte
g*~a
the mouth of the latrine and allowed to extend about 6 inches
above the mouth is a commonly used method. Clay is packed
around the protruding keg, the
slab is placed on top, and then
the superstructure is built.
To prevent caving in areas
where the walls are likely to
cave in the entire depth of the
latrine, plaited bamboo or
wickerwork linings, or cylinders
made of cement, clay, wire,
wood, or sheet metal are used.
Drums or kegs placed end on
end have been frequently used
in the Philippines. The bamboo
linings are very satisfactory es-
pecially if coated with coal tar
or some other wood preserva-
tive. In some places the latrines
are bored to a depth of 12 or
14 feet below the water level.
Bamboo under water lasts years
without a preservative; there-
fore, in these areas it is not
necessary to use a preservative
on the portion of the lining that
will remain under water. The
wood extending above the water
level is more likely to rot or be
eaten by insects. In many trop-
ical countries the bamboo will
last as long as the latrine and
is cheaper to replace if neces-
sary than to use preservatives.
A woven bamboo lining is shown
in fig. 41, a, and an enlarged
sketch showing the weaving is
shown in fig. 41, 6. These woven
linings were first used in Java
and have been used satisfactorily in the Straits Settlements, and
are extensively used in the Philippine Islands and other coun-
tries. The bamboo cylinder can also be made by tying long
Fig. 41. Woven-bamboo cylinder partly
made. This makes a very satisfactory
latrine lining in most places. The sec-
tion a is a temporary support to keep
the ribs straight while weaving; & is an
enlarged section of the woven-bamboo
latrine lining.
46,4
Yeager: Bored-hole Latrines
725
bamboo strips to hoops. In some places where they do not have
bamboo they fasten strips of another kind of wood to metal or
wooden hoops.
Dr. Victor G. Heiser suggested the use of an open mesh gal-
vanized wire screen as this would allow the bacteria naturally
in the soil to act on the dejecta and also allow permeation of
the latrine contents into the soil. We have recently tried wire
screen made into cylinders as shown in fig. 42, a. The only
disadvantage is that chemical action might cause rapid disinte-
gration of the metal. In some places the bamboo cylinders
would probably last longer, but wire can be obtained in places
where there is no bamboo. A thin wire netting ordinarily called
chicken wire net can be reenforced with wood or iron hoops,
but is too flimsy for practical use. A mesh of i-, |- or 1-inch
heavy wire does not require the hoops and withstands corrosion
longer. These cylinders cost about 3 pesos for each section 3
feet long. Sheet-metal cylinders can be made as described in the
next paragraph, but numerous holes should be chopped into the
metal to allow better action on the dejecta.
Silting-earth reenf or cement. — In order to prevent caving in
silting earth a solid lining is required. Bamboo cylinders are
very satisfactory in some places, but we have frequently found
Wooden on me fa/ strips*
^Oi/ f drum
Fig. 42. Various latrine linings, a, A galvanized-iron wire-net lining which can be used
in places where white ants eat the bamboo or where bamboo is not available; b, stove-
pipe double wall, "wall method" of making a lining. In ordinary soils numerous holes
can be cut into the metal to allow better action. In silting soils the sections are
added one at a time as boring proceeds. A few dents hold the cylinders together;
c, four to six empty metal oil or tar drums placed end on end are frequently used as
linings for latrines. The heads of the drums are cut out with a chisel. Wooden or
metal strips are nailed to the drums to make strong joints. These are often used in
silting sand.
726 The Philippine Journal of Science 1931
it necessary to cover the cylinder with a thin coat of clay or
weak mixture of cement in order to prevent very soft sand from
silting through the small openings in the cylinders.
In most places in the Philippines we use cylinders that we can
get for little or nothing. Heavy sheet-iron oil drums with the
ends cut out and tar and cement drums have been very satis-
factory. We frequently use wooden cement kegs placed end
on end.
When using sheet-metal linings in soft sand we rivet the cy-
linders with five or six small rivets. We make the cylinders 3
feet long for convenience in handling. A few slits can be
chopped into the lower cylinders to allow a rise and fall of water
if necessary. Each cylinder is slit in four places at one end to
allow it to be inserted an inch into the next cylinder.
Stove-pipe method of lining. — When only very thin sheet
metal is available we use the stove-pipe-well method. A number
of cylinders are made in two diameters each 3 feet long so that
the smaller cylinders can be telescoped or slid into the larger
cylinders. These make an excellent reenforcement and are
convenient for boring in very soft sand. The joints overlap
in the center and a few dents with a hammer prevent slipping.
Fig. 42, b, shows the position of the joints.
If the soil is stiff enough to hold its shape long enough to bore
to the required depth, the lining is put down in one or two long
pieces. If the soil constantly caves in from the sides while bor-
ing, the cylinders, which are a couple of inches larger in dia-
meter than the auger, are put down the latrine one section at
a time, and the sand is pulled up through the cylinder by the
auger. The cylinders are pushed down every few minutes in
order to block off the silting soil as rapidly as the auger cuts.
If thin metal cylinders are used, a hoop of iron should be fast-
ened around the bottom cylinder to maintain a circular opening.
If this is not done the sides will be compressed by the mud or
sand and the auger will not turn or will hook under the edge
of the cylinder.
Iron oil drums with the ends cut out and placed end on end are
used in the same way as the sheet-metal cylinders, and have
been very useful in reducing the cost of latrines in soft silting
sand and mud near Manila. A satisfactory way to joint the
cylinders together is shown in fig. 42, c. Three or four strips
of wood hold the joint solidly. Nails are driven through the
wooden strips and drum and clinched on the inside. Nails are
46,4 Y eager: Bored-hole Latrines 727
difficult to drive into heavy oil drums unless holes are previously
punched through.
Slabs or floors. — Under exceptional circumstances we have
allowed the use of wooden slabs or floors in latrines but for ob-
vious reasons usually insist upon the use of cement slabs. A
variety of slabs have been designed in many countries with as
many different sizes and shapes of holes. In some places plain
reenforced concrete slabs are made with rectangular holes in
the center. In other places they use elevated treads or so-called
foot plates for the feet to prevent fouling. A slab that has been
used successfully in Java is shown in Plate 7, fig. 2. Plate 1 is
one of the types recommended by the Government of Madras,
India. They also use a circular slab. The Java slab costs less
and is the smallest slab we have heard of. In other places they
use slabs over 4 feet long. In the Philippines we most fre-
quently use the reenforced slab shown in Plate 2. This slab is
30 inches wide and 36 inches long. It is 2\ inches thick at the
outside with a sloping surface for drainage reducing the thick-
ness to 2 inches at the edges of the holes. The edges of the
holes are cut back to prevent fouling. Knowing of two in-
stances in which children fell through holes 8 by 18 inches in
one country, the rectangular hole in the slab used in the Philip-
pines is made only 5f inches wide and 13 inches long. In cast-
ing the slabs a notch 3 inches on each side is made at each corner
for the posts of the superstructure. This allows the walls to
be built close up to the edge of the slab.
Another kind of slab is also used in the Philippines ; it is of
the same general dimensions but with the hole narrowed at the
front and with two elevated treads or foot plates for the feet.
This slab costs a little more to make than the plain slab, which
costs from 1.85 to 2.45 pesos including J-inch twisted wire re-
enforcement, a 1, 2, 3 mixture of concrete, and the labor. The
plain slabs are more easily handled in shipping than those with
treads, and up to the present time have not been found fouled
any more frequently than the slabs with foot plates.
Dr. W. P. Jacocks designed an excellent latrine floor or "squat-
ting plate" for Ceylon. This plate is made of 18-gauge pressed
steel and can be purchased for 4 rupees or less from Messrs.
Walker, Sons & Co., Ltd., Colombo (fig. 43).
The floors of latrines should always be at least 18 inches above
the highest water level. If the water level is several feet below
the surface of the earth and the rainy or flood season does not
728
The Philippine Journal of Science
Elevation
1931
3 18 W. G. Plate
%: I" holes
Fig. 43.
A steel latrine floor, or squatting plate, designed by Dr. W. P. Jacocks for
in Ceylon. The cost is about 4 rupees, about 2.60 pesos.
raise the water to the surface the slab can be placed directly
over the hole on a level with the surrounding soil.
In places where the water floods over the area the floor and
the superstructure can be built upon mounds of clay. In these
places the lining can be allowed to extend above the surface of
the earth to the required level and clay, bricks, stones, cement,
or other material can be placed around the protruding lining to
support the slab. We sometimes use blocks of tuff or adobe ce-
mented with clay or lime, but in most instances on account of the
possibility of spreading infection from the latrine we use cement
to make the structure solid and without crevices.
At times we run the cement a foot or more below the surface
down around the lining or dig a small trench about a foot away
48' 4 Y eager: Bored-hole Latrines 729
from the bored hole and fill this with cement. While there is
probably no seepage under a heavy slab with its support, the
above precaution tends more completely to block any exit of in-
fection. In places where the people do not have enough money
to build a solid structure, we frequently use discarded oil drums
with the ends cut out for the support of the slab. The drum
extends about a foot into the mouth of the latrine and 2 feet
above the surface where the water floods the area to about a
foot in depth.
The water trap.— There are many houses equipped with flush
water closets emptying into bored latrines. In some of these
two or three bored holes are connected together with pipes and
are better than many septic tanks. Where they do not have
piped-in water but have the money to purchase a porcelain water
closet, the bowl is placed on the slab directly over the bored hole
and after using is flushed with a bucket of water.
One of the advantages of the bored-hole latrine installed in
suitable places where the subsoil water is encountered at a depth
of about 16 feet, is that mosquitoes, flies, and other objectionable
insects do not breed in the latrines and no bad odors are emitted.
One objection to the bored latrine in places where the ground
water lies within a few feet of the surface of the earth is that
it furnishes a breeding place for flies and mosquitoes and is of-
fensive unless properly constructed. The ordinary slab is not
satisfactory in these places and the cost of vitreous china bowls
is too great for a large proportion of the rural inhabitants. To
meet the demand reenforced concrete water traps have been
made. The design first used was copied from a porcelain water
closet. A small bucket of water is sufficient to flush the bowl
completely. One feature of the trap is a clean-out hole, which
facilitates cleaning when necessary. These traps have been
greatly improved by making them longer and putting foot treads
on them. We have now two types of traps so designed that the
dejecta must fall where supposed to at the back of the water
closet. It is impossible to squat on these water closets back-
wards because the treads slope to the front and throw a person
off balance when he tries to squat on the trap the wrong way.
One of these water closets is designed to be placed on top of a
concrete slab, and the other is designed so that it fits flush with
the slab, except the treads, so that the floor drains into the trap.
The traps are made of cement and cost about 4 pesos each for
264209 13
730
The Philippine Journal of Science
1931
. Boor of .
supemfrocwtt
fioredfafe
~&perjtrttct(ff*e
Water hap
a.
Fig. 44. Latrine and water closet, a, The bored-hole latrine complete with superstructure.
The lining and cement casing are not used in soil that does not cave into the latrine.
Metal drums are usually used instead of cement casings where the water rises to the
surface; bt a cement water closet that can be flushed with a bucket of water. About
two hundred of these water traps are giving excellent service in the Philippines. The
sloping foot rests make the user sit on the water closet correctly. The rests throw one off
balance if he attempts to squat backwards. These traps cost about 4 pesos each. They
absolutely eliminate fly and mosquito breeding and foul odors.
material and labor. A few baked-clay traps have been made,
but these are not as satisfactory as the cement water closets
(Plate 4 and figs. 44 to 46).
We recommend these flush water closets for private families
only and not for use in public latrines, because careless people
block them with rubbish. Nearly two hundred water traps have
been in use over a year and are a very satisfactory improvement
to latrines where the water level is high.
These traps have been placed over latrines that were covered
with swarming maggots and emitted very objectionable odors.
46,4
Y eager: Bored-hole Latrines
731
ol*"
3"
■"" r
5"
*• & U »
1
,
1
//
%i
(
(
4J ^
\
^
1
*t
} •
55
1
3
,1
*
1 , y
7
k^ ,
f
(
\ \ J ! c
\
1
*
^
t
V,
to match with
ihose on the
water trap.
)
'
>
*"* 1
f
1
.#/ Cement
L-A"£j( /fiwsh h3 &
11/ 1:3-6 concrete
<>4
t
1 m" J> 1
Fig. 45. Reenforced concrete slab for use with water trap. Elevated type.
After installation of the traps and cleaning the latrines there
was no longer any fly or mosquito breeding and the odor dis-
appeared immediately. Another advantage of the trap is that
the people learn almost immediately to use water or paper in-
stead of sticks and other articles, and the latrines will serve a
greater length of time.
Superstructures. — In this section on latrine construction there
is no attempt to discuss the building of superstructures. There
are so many kinds of superstructures that are satisfactory and
the details are of such little importance, as far as the prevention
of disease is concerned, except in one point, that it is unneces-
732
The Philippine Journal of Science
1931
PlO. 46. Latrine equipped with water trap sunk flush with the slah. This trap costs only
8 pesos. Note the sloping foot rests and the small vent, which can be opened to relieve
pressure in an air-tight latrine. The washings from the trap fall from the lip and do
not follow the trap down to the wall of the latrine.
sary to take the space required to include a description in this
article. Sometimes the latrines are bored so that the seat or
slab can be placed in the house or in a position easily accessible
46' 4 Y eager: Bored-hole Latrines 733
by an elevated board walk leading from the house to the latrine.
There are many places in our rural areas where they do not build
a complete superstructure, but only palm leave partitions on three
sides of the latrine with a screen of palm leaves erected a couple
of feet in front of the entrance for privacy. There is only one
detail in the building of superstructures in which we are partic-
ular and in which the builders of the superstructures are more
or less careless. In some places they build the superstructure
so large that a space of a foot or more of earth is left uncovered
by flooring between the slab and the walls of the superstructure.
In these instances there is danger of soil pollution around the
sides of the slab where the earth is shady and moist and this
furnishes an excellent culture medium for parasitic larvae and
other disease organisms. We insist upon the walls of the build-
ing being placed against the slab so that the floor of the latrine
is all concrete.
A complete bored latrine of the usual type is shown in fig. 44,
a. The bamboo lining and a cement or iron casing shown in
this picture are not needed in solid ground which does not cave
in.
The laborers' privy as built with a seat by the United Fruit
Company is shown in Plate 5. This sketch also shows their me-
thod of lining latrines.
Dr. Louis Schapiro forwarded a pamphlet to us from Siam
showing an ingenuous latrine that is fool proof as far as keeping
the hole in the floor covered. In order to use this latrine the
lid must be pushed back. When the lid is in the open position
the door cannot be opened because it strikes a handle on the lid.
Therefore, in order to leave the latrine the lid must be closed so
that the door can be opened (Plate 6).
BACTERIAL MIGRATION
Regardless of the results of the experimental work attempting
to determine the limits of bacterial migration, it is advisable to
frequently examine well water for pollution in every community.
Some of the work done in the past throws some light on the pos-
sibilities of pollution. There is considerable evidence showing
that bacteria do not travel more than a few feet in dry soil and
in some instances not even one foot from the latrine, but more
information is needed to show the distance bacteria will travel
in ground water. Bacteria have been carried several thousand
feet in ground water flowing through fissures in lime stone and
other formations.
734 The Philippine Journal of Science 1931
W. A. Hardenbergh discusses the results of a considerable
amount of work on pollution in his book on home sewage dis-
posal. The United States Public Health Service, through an ex-
perimental board directed by Dr. Charles Wardell Stiles, reports
bacterial migration 200 or 300 feet, and states that possibly-
greater distances are covered under most favorable conditions.
Stiles, in "The principles underlying the movement of Bacillus
coli in ground water, with resulting pollution of wells," states
that B. coli were found in ground water 65 feet from a polluted
trench. These bacteria evidently traveled through a fine sand
with an effective size of 0.13 millimeter. It took 187 days for
B. coli to travel this distance, and the migration was only in the
direction of the flow of the ground water. In another article,
"Experimental distribution of Bacillus coli in the soil under and
near pits," by C. W. Stiles and C. L. Pfau, it is stated that B. coli
were carried in ground water up to at least 232 feet.
Space cannot be given in this article to discuss the viability of
organisms, but the resistance of the pathogenic bacteria which
are likely to infect man through drinking water is not great.
Most of these bacteria die within a few days. Typhoid bacilli
might live months in polluted soil on the surface, but the bac-
tericidal action at ground-water levels is greater. Kligler
states, typhoid and dysentery bacilli may be recovered up to
seventy days in moist natural soil. The rapidity of time in
transmission is an important factor in well pollution.
Although Stiles found no convincing evidence of the travel of
B. coli against the flow of the ground water, there is no reason
why these bacteria and other organisms could not migrate
against the flow in some places. In areas where the water does
not flow constantly in one direction or at times is almost at a
standstill, the motility of the organisms as well as migration by
growth would be factors in the spread. Cholera and choleralike
vibrios can actually travel against the flow of a current. Schobl
has demonstrated this point many times. Hardenbergh states
that "pollution appears to travel against the flow of ground
water as well as with it." Other workers have made similar
observations. However, the migration of bacteria against the
flow is relatively unimportant.
A factor in migration that is sometimes not considered is the
kind of soil in which small channels form. In some soils there is
practically an even filtration. In other soils of the same effective
size of sand there is a cohesive quality which affects filtration.
An even filtration might be expected, but examination reveals
4®, 4 Y eager: Bored-hole Latrines 735
that the water is flowing into the well in small rapid-flowing
streams through a few long channels some of which are branched.
In these places the bacteria are not subjected to the same filter-
ing action as in a soil where there is even filtration.
Although Israel J. Kligler 2 states that the "pollution of wells
is usually surface in origin/' the direct pollution through ground
water is pointed out in his summary, as follows:
The problem has been approached both from the experimental and
practical standpoint. In the laboratory repeated tests have been made to
determine: (1) the viability of the typhoid and dysentery bacilli in soil
and in excrement under different conditions; (2) their ability to penetrate
through columns of soil of different porosity; (3) their viability in septic
fluids and effluents; and (4) the nature of the antagonistic factors in
soil and septic material which influence the viability of these microorgan-
isms. In the field work various types of privies of different ages were
examined particularly with regard to (1) the extent of pollution of the
soil surrounding these privies; (2) their relation to well pollution; and
(3) the passage of material from the privies through the soil to adjoining
wells.
The main conclusion arrived at on the basis of both the experimental
and field observations is that in moderately compact clay, sand-clay, or
sandy soil, free from cracks, the possibility of subsoil pollution of the
ground-water is negligible, provided the ground-water level is more than
ten feet below the polluted area.
The following facts were established :
1. The typhoid and dysentery bacilli succumb rapidly on exposure to
an unnatural environment, (a) Both typhoid and dysentery bacilli die
out in 1 to 5 days in septic tanks. (6) In solid feces the typhoid bacilli
may survive for a period of 10 to 15 days, while the dysentery bacilli
rarely survive longer than 5 days. The paratyphoid bacilli are the most
resistant members of the group; the Shiga dysentery bacillus is the most
sensitive, (c) The survival period of these organisms in soil is greater
than in either feces or septic fluids, and varies particularly with the mois-
ture and reaction of the soil. Temperature effects the viability, but the
two main factors normally are moisture and reaction. In moist natural
soil of a pH value of 6.6-7.4, the typhoid and dysentery bacilli may be
recovered up to 70 days. In the same soil dry, the bacilli are not recov-
ered after 2 weeks. In moist acid soils, pH 4.8-5.4, 90 per cent of the
inoculated bacilli die out within the first 10 days, the others may survive
as long as 30 days. All the organisms survive longer near freezing temper-
ature (4° C.) than at higher ones (20-37° C). (d) The antagonistic ac-
tion of soil bacteria on typhoid and dysentery bacilli is due largely to the
alkaline reaction resulting from their metabolism. Specific inhibitive subs-
tances are, however, elaborated by some soil bacteria, notably Bacillus
fluorescens and Bacillus proteus.
2 Rockefeller Institute Monograph 15.
736 The Philippine Journal of Science 1931
2. The spread of pollution from a focal point is limited in scope, (a)
Typhoid and dysentery bacilli under experimental conditions were not
observed to spread laterally to any appreciable extent, although they were
carried vertically through a column of 2 feet of porous soil. In denser
soil they failed to penetrate through 1 foot. (6) In the field, where the
subsoil was free from pollution, either near pit privies or near tile pipes
from septic tanks, contamination extended downward to a depth of 5 to
3 feet, and laterally only about 3 feet, from the bottom of the pit or tiles.
(c) Heavy rains or constant dripping of water may carry surface pollu-
tion to a depth of 10 feet.
3. Pollution of wells is usually surface in origin, (a) There was no
correlation between the type or proximity of the privy to the degree of
contamination of the adjacent wells. The purity of the well water varied
rather with the condition of the well. Driven shallow wells with pumps
were, as a rule, free from contamination, while dug wells with pumps
or buckets were generally grossly polluted. (6) Experiments with fluores-
cine failed to show subsoil pollution of wells from privies, but proved
in some instances at least the possibility of surface contamination.
According to Kligler, F. A. A. F. Eykin and G. Grijns, working
in the Tropics, made similar observations. They found very
little pollution in the soil around pits not reaching the ground-
water level and in only one case traced pollution 5 meters from
the pit. Bacillus coli was not found at a depth of 20 inches
from the bottom of the pit, but during wet weather the pene-
tration was about three times as great. As stated by W. A.
Herdenbergh —
In the case of high ground water, these authorities seem to think that
much pollution is from the soil directly into the ground water and thence
to the well, with the privy having no part in the process. This would ap-
pear to fall under the head of surface pollution and illustrates how me-
chanical (or animal) transportation of pollution may be an important
factor in the spread of disease.
The Commission on Additional Water Supply for New York
City found that polluted water was rendered safe to use after
flowing through 25 feet of fine sand. In abstracting a report of
this observation W. A. Hardenbergh states :
The report of the Commission on Additional Water Supply for New
York City, made by Burr, Hering and Freeman, records some experiments
on the same subject. The tests were made at Elmont, L. I. While it is
stated that the passage of polluted water at low velocities through twenty-
five feet of the finer sands, such as are found in Long Island, will render
the water safe to use, it is also shown that, under severe conditions sewage
bacteria and B. coli may pass through soil for a considerable distance.
In most cases, a lesser distance than twenty-five feet may be considered
safe, it is stated.
46,4 Y eager: Bored-hole Latrines 737
Shallow pits if unprotected from surface water, according to
the investigations of the Public Health Service at Wilmington,
North Carolina, and other observations, are a source of danger.
In the work at North Carolina, pits filled with surface or rain
water carried pollution to the ground water below, and from pits
reaching the ground water B. coli traveled a distance of 200 feet.
Most investigators agree that if the ground water is not pol-
luted the chance of infecting wells through the soil is very re-
mote, and that most water supplies are infected from surface
pollution. Quoting Hardenbergh :
Dr. C. T. Nesbitt, at that time Health Officer of New Hanover County,
N. C, made a series of tests in 1917 at some mill villages near Wilmington.
He found pollution of the ground water from effluents from septic closets,
in some cases up to twenty feet from the effluent pipe, beyond which
distance he made no tests. The results in this case were generally such
as to indicate that the sand-clay soil of that region does not fully protect
the ground water from fecal pollution, nor prevent the travel with under-
ground water of such pollution for uncertain distances.
This indication was borne out by the bacteriological examination of
about 700 shallow driven wells located in the city and county. The only
wells of this kind free from pollution were those located 200 to 500 yards
away from any concentrated source of pollution, as stables or privies.
The bacterial counts in those wells not so located were extremely high, and
the presumptive tests for B. coli were almost unfailingly positive.
I do not have a reprint of the work on these 700 shallow wells,
and do not know if there was surface pollution or not, but in
many other places a careful examination has revealed surface
pollution of most of the wells.
In Manila there are 42 artesian wells from 345 to 900 feet in
depth, which are not likely to be contaminated from ground
water. Pollution was found in many of these wells, but careful
examination showed the possibility of surface pollution. After
the concrete platforms around these wells were elevated and
the well heads repaired presumptive B. coli were seldom found
and the bacterial counts were reduced to satisfactory limits.
Shallow wells would not have shown such good results. In fact
satisfactory water drawn from a depth of 20 feet has never been
found in Manila. We have no evidence showing that bored-hole
latrines reaching the ground-water level have ever infected a
properly constructed deep well. In many villages the cost of
numerous shallow wells, which are sources of danger regardless
of bored-hole latrines, is greater than the cost of a deep well;
738 The Philippine Journal of Science 1931
therefore deep wells are recommended where satisfactory piped-
in water cannot be obtained.
Rosenau * shows a picture of the "popular idea of how wells
become infected from surface pollution," and states, "this rarely
takes place in rural districts, as the soil can usually hold back
most of the impurities." Similar pictures are shown in many
elementary books on hygiene, public-health pamphlets, and pos-
ters. It is more reasonable to show direct pollution from the
surface where bacteria obviously have easy access to the well.
According to Rosenau, "The viability of typhoid bacilli in
feces is very variable, depending on the composition of the feces
and the varieties of other bacteria present." Sometimes typhoid
bacilli in faeces perish in a few hours, and under other condi-
tions have been found to live five to seven months. The life
of the organism in privies and in water is usually comparatively
short. "In nature they seldom, if ever, live in water beyond 7
days, and are often dead in 48 hours." They probably live
longer in clean water than in contaminated water, but soil pol-
luted on the surface is most dangerous. The deleterious effect
of antibiosis, chemicals, temperature, light, dissociation, sun-
light, filtration through soil, and other factors affect bacteria.
Rosenau refers to the use of dyes and chemicals to determine
the sources of pollution. He regards these tests as valuable in
indicating the possibility of danger under certain circumstances
and finds them useful in discovering the sources of pollution
near wells or in limestone formations. He points out the pos-
sibility of error in concluding that microorganisms and danger-
ous pollution travel an equal distance to the chemicals, stating
"the soil has well-known filtering power when free from fissures
or open channels and is capable of removing bacteria and oxi-
dizing large quantities of organic matter."
On the other hand Rosenau records the travel of Bacillus
prodigiosus to a distance of 200 meters in forty-two hours. He
does not describe the soil nor state whether or not there was
surface travel. The cultures were poured into the ground.
Rosenau cites many examinations under, "interpretation of
sanitary water analyses," in some of which there was evidently
pollution from soil surcharged with organic matter, and another
instance in which there was remote pollution in which organic
matter was mineralized and the bacteria held back by the soil,
* Preventive Medicine and Hygiene, page 951.
46» * Y eager: Bored-hole Latrines 739
and a number of wells in which there was direct infection from
the surface.
It is believed by many that the B. coli and count tests do not
give the information required. Some pathogenic bacteria will
die where B. coli might survive in an acid soil. The cholera
vibrio lives best under alkaline conditions. Dr. Otto Schobl,
of the Bureau of Science, Manila, suggested the use of a cholera-
like vibrio which is easier to identify than B. coli and responds
to enrichment in peptone broth. This organism is a much better
indicator for cholera than B. coli, and work is now under way
in the Philippines on the migration problem. Doctors Ramirez
and Basaca, of the Bureau of Science, have given much of their
time to the bacteriological technic.
In an article, entitled "Well pollution and safe sites for bored-
hole latrines," an attempt was made to caution health officers
as to the possible danger of installing latrines near shallow wells.
This piece of work was not intended as a scientific check on
previous bacteriological work on migration that has been pub-
lished by a number of bacteriologists. The work had to be
done in a very limited length of time, and the number of ex-
aminations were limited. Latrines were being installed within
a few feet of shallow wells, and the experiment showed that
bacteria in that area would travel in subsoil water considerably
farther than this distance. As stated in the article, "The en-
tire experiment was of such short duration that we make no
pretence of presenting a complete piece of work. However, the
results give a fair indication of the degree of possible contami-
nation of water-supply situated within short distances of bored-
hole latrines in a similar soil." This was only a preliminary
piece of work, and it served its purpose until more complete work
could be undertaken.
Credit should be given in this article to all persons who have
contributed toward the development of the bored-hole latrine,
but the names of individuals other than those mentioned are
not available.
The three following paragraphs are quoted from a report
submitted by Dr. John L. Hydrick to the Rockefeller Founda-
tion:
lAfter the rural hygiene campaigns in Java had been in operation for a
few months, the sanitary inspectors noticed that in the areas in which
the simple pit latrines were deep enough to reach groundwater the floors
of the pits were covered with black sludge and a thick scum floated on
the water.
740 The Philippine Journal of Science mi
Since the odors of fermentation from these latrines were not more
objectionable than those from dry pits, Dr. van Breemen, city health of-
ficer of Batavia, decided to carry out some experiments with a type of
latrine devised several years ago by one of the field officers of the Dienst
der Volksgezondheid in Nederlandsch Indie.
This field officer had noticed that the small borers used on the estates
for digging holes for fence posts and telephone poles were easily han-
dled by the laborers and that with very little difficulty holes could be
dug to the groundwater level. He suggested that these borers be used
to make simple pit latrines, since fecal material deposited in a deep pit
which reaches groundwater would undergo fermentation. The action would
be similar to that of a septic tank; a narrow deep hole should be usable
over a long period of time, and, on account of its small diameter, its
walls would be less liable to cave in after heavy rains.
There are published articles on latrine boring that have not
yet been received. This publication covers methods developed
locally, descriptions furnished by manufacturers, and reports
forwarded by Dr. Van Wesep and Mr. Rollin C. Dean, of the
Rockefeller Foundation. Dr. Victor G. Heiser has contributed
much information and many valuable suggestions at frequent in-
tervals. Many officers in the Straits Settlements and in the
Philippine Health Service, especially Dr. Jacobo Fajardo, director
of health, and Dr. Gabriel Intengan have given active coopera-
tion in the work. Mr. Cecilio Marcelino has given much of
his time in making molds for water traps. We are indebted
to Mr. Eugenio Viana, superintendent of San Lazaro Hospital,
for his active cooperation in the construction of many prelim-
inary designs of cement slabs, water traps, and other work.
Mr. Manosa, chief of the Sanitary Engineering Department
of the Philippines, and Messrs. Diaz, Claustro, and Bagabaldo
have given valuable service in latrine installation.
I am indebted to Dr. William H. Brown, director of the Bu-
reau of Science, Philippine Islands, for editorial suggestions and
corrections. Mr. R. C. McGregor, associate editor of the Jour-
nal of Science, has rendered valuable aid in editorial corrections
of this article; and Macario Ligaya and Francisco Rafael, of
the Bureau of Science, have spent a great deal of time redraw-
ing many of the pictures.
ADDRESSES OP MANUFACTURERS AND DEALERS AND WHAT THEY SELL
There are many manufacturers and distributors of boring
equipment, but the following list will be sufficient to allow a
selection of inexpensive equipment as well as modern up-to-date
machinery.
46, 4 Y eager: Bored-hole Latrines 741
The National Supply Corporation, 120 Broadway, New York City, and
185 Queen Victoria Street, London, E. C. 4., have in stock or can obtain
almost anything in boring equipment known.
The Oil Well Engineering Co., Ltd., Cheadle Heath, Stockport, England,
and R. Richards & Co., Upper Ground Street, London, S. E., deal in an
extensive line of boring apparatus.
Werf Conrad, Drilling Outfits Department, Haarlem, Holland, manufac-
ture the Banka hand drill, which is not practical for bored-hole latrine
work, but they have derricks, a variety of augers, chain tongs, and many
useful accessories.
The Ingersoll-Rand Company, 11 Broadway, New York, manufacture
an enormous variety of power-driven machinery that will drill anything.
They sell the "calyx" hand-power geared outfit that can be equipped with
a clay auger to bore 16-inch holes. This apparatus sells for about 1,750
dollars. They also sell drills for making blast holes in rock.
The Keystone Drill Company, Beaver Falls, Pennsylvania, with offces
at 170 Broadway, New York, manufacture a large variety of power drills,
and numerous types of clay augers, sand pumps, bailers, rock drills, and
other equipment.
The Star Drilling Machine Company, Akron, Ohio, manufacture port-
able well-drilling machinery, confined to the churn or percussion-type drill,
which is not practical for latrine boring. They manufacture many types
of auger bits and accessories.
The Okell-Well Machinery Corporation, 2035 Bay Street, Los Angeles,
California, quote a price of 1,798.91 dollars for a complete boring ma-
chine including a seven-horse-power motor and a full line of accessories.
R. R. Howell & Co., Minneapolis, Minnesota, are manufacturers and
jobbers of a full line of drilling machinery, augers, and supplies.
Sweeney and Gray Co., Inc., 81 Sixth Street, Long Island City, New
York, manufacture boring equipment, and sell a clay auger with a hinged
bottom for about 100 dollars. The shafting costs 40 dollars.
The Specialty Device Co., 106 West Third Street, Cincinnati, Ohio, man-
ufacture the "Standard" earth auger discussed in this paper, a quadruple
expansion brace for holding the shaft, and other useful accessories. These
products are sold through the A. J. Alsdorf Corporation, 330 South Frank-
lin Street, Chicago. The No. 10 auger equipped with extension blades
costs 6.60 dollars. The brace costs 4 dollars. A 25-foot coupled shaft
costs 3.75 dollars. Couplings and bolts cost 2 dollars.
Lang-London-Ltd., 34 Gray's Inn Road, Holbom, London, W. C. L., sells
the Lang earth borer discussed in this paper.
Iwan Brothers, South Bend, Indiana, manufacture the Iwan post-hole
auger, disk auger, and sand-digging tools. These augers can be purchased
from the National Supply Co., 185 Queen Victoria Street, London, and from
Lindeteves-Stokvis, Amsterdam, Holland; Batavia, Java; and Penang,
Straits Settlements.
The Budda Company, Harvey, Illinois, with an export office at 30 Church
Street, New York, manufacture the Budda-Hubron machine. The trailer-
type machine for boring holes 20 feet deep costs 2,275 dollars and the
truck-mounted type costs 2,125 dollars. This is the most compact power-
driven machine on the market.
742 The Philippine Journal of Science 1931
The Gus Pech Foundry and Mfg. Co., 200 Second Avenue, S. W., Le
Mars, Iowa, manufacture a practical machine-driven auger, the No. 2
Monitor mounted boring machine, and a variety of auger bits, stone hooks,
and other accessories that are valuable in latrine construction.
The Howells Mining Drill Co., Plymouth, Pennsylvania, manufacture
the twist drill illustrated in this article, which has been especially adapt-
ed to our work for drilling blast holes in adobe, shale, and other hard
formations. They also make the hand geared machine known as Howells
prospector's drill, which should be equipped with bits to bore 1-inch by 2i
foot blast holes at variable depths up to 18 feet, and the weighted tripod
regularly furnished or the stand used on their Spry Little Giant slate
drill. They also make air and electric drills.
Oilwell Supply Co., London, and 215 Water Street, Pittsburg, Penn-
sylvania, sell all kinds of boring apparatus.
Armstrong Mfg. Co., Waterloo, Iowa, sell a large variety of drills.
McKieman-Terry Drill Co., 115 Broadway, New York, sell many kinds
of drilling apparatus.
EQUIPMENT AND WHERE TO OBTAIN IT
Dollars
United States
currency.
Iwan post-hole auger bottom, 14-inch (dozen) 67.20
Iwan post-hole auger bottom, 16-inch (dozen) 81.60
Iwan post-hole aug^er bottom, 16-inch, extra heavy
blades (each) 12.50
Iwan Bros., South Bend, Indiana.
National Supply Corporation, England.
Lindeteves-Stokvis, Amsterdam, Holland; Batavia,
Java; and Penang, Straits Settlements.
Chain tongs, 2i-inch Vulcan bijaw (each) 3.50
J. H. Williams and Co., Brooklyn, New York. Na-
tional Supply Corporation, New York and Eng-
land.
Nearly all local hardware stores.
Pulleys (each) 100
Local stores.
Wilson pipe wrench (each) 15.00
National Supply Corporation.
Drop-bottom double-bit augers for earth, sand and gra-
vel, 16-inch (each) 45.00
Gus Pech Manufacturing Company.
National Supply Corporation.
Okell-Well Machine Corporation.
R. R. Howell and Co.
Sweeny and Gray Company.
Clay auger, 151-inch, No. 1, with 20-foot boring rods
(each) 95.00
R. Richards and Co., London.
Clay auger, 14- and 16-inch drop bottom.
R. R. Howell and Co.
Gus Pech Manufacturing Company.
46' 4 Y eager: Bored-hole Latrines 743
Dollars
United States
Lang earth borer, 14-inch, complete with sand-boring at- Cm'nC7'
tachment and accessories (each) 9g#00
Lang-London, Ltd., England.
"Standard" auger No. 10, 10-inch, with cutters complete
with 20 feet of shaft (each) 10 35
A. J. Alsdorf Corporation.
The Gus Pech or No. 2 Monitor boring machine (each) 842.50
Gas Pech Manufacturing Company.
National Supply Corporation.
Okell-Well Machine Corporation.
The Budda-Hubron machine (each) 2 275.00
The Budda Company, Harvey, Illinois.
Hand-twisted drills, geared drills, electric- and air-dri-
ven drills for making blast holes in shale, slate, adobe
rock, laterite, and other hard formations.
Howells Mining Drill Works, Plymouth, Pa.
Machine-driven rock drills for making blast holes in
hard rock.
Ingersoll-Rand Company.
Drill steel for making bars is sold by local dealers and by Ingersoll
Rand who carry in stock a large supply of standard sizes and shapes
such as hexagon, round, square, pentagon, cruciform solid, and twisted
concave. They also have finished sets of drill steel.
SUMMARY
The Iwan post-hole auger with locally made shaft and turning
handles, an inverted V hoisting frame equipped with a compound
pulley and a shaft supporting brace or hinged-door platform is
recommended as the cheapest fast-cutting equipment for boring
latrines in all soils except hard formations, silting sand, and
soft mud.
Various shafts, turning handles, braces, hoisting equipment,
and miscellaneous accessories are discussed.
Useful augers for general boring including very soft soil are
the locally made augers, the auger used by the United Fruit
Co., and a large number of augers on the market.
Useful augers for boring in very soft soil and silting sand
are the locally made augers with hinged blades, the augers used
by the United Fruit Co., the Iwan augers equipped with locally
made valves, the Lang augers, and a variety of bailers, pumps,
and other augers regularly supplied by dealers.
A number of hinged shafts to facilitate dumping are shown.
The method of preventing caving of the latrine walls while
boring in mud and quicksand is described.
A rapid practical method of installing latrines in tuff, or so-
called adobe rock, and other very hard formations is described.
744 The Philippine Journal of Science
The method described has been used successfully since October,
1929.
Power-driven drilling machines are recommended for rapid
economical boring where large numbers of latrines are to be
installed and there is sufficient space for the operation of the
machines.
Linings to prevent caving of the walls of bored latrines and
the methods of using linings in silting formations are described.
Water traps made locally of baked clay or cement, which
greatly improve the bored-hole latrine, especially where the
ground-water level is near the surface, have been used success-
fully. These traps are described and illustrated.
A variety of cement slabs and a metal latrine floor are
recommended.
The details of building superstructures are not given, but
the importance of constructing the slabs and walls so that no
uncovered earth is exposed to contamination is emphasized.
A brief description of bacterial migration is useful in locating
safe sites for bored-hole latrines.
Dr. Victor G. Heiser suggests emphasis on proper supervision
so that bored latrines are not installed in places where there
is danger of infecting domestic water supplies.
A list of manufacturers and dealers with their addresses and
the materials they sell that are useful for making bored-hole
latrines is included.
A list of bored-hole latrine boring equipment, where to ob-
tain it, and the prices are given.
BIBLIOGRAPHY
Rosenau, M. J. Preventive Medicine and Hygiene (1927).
Hardenbergh, Wm. A. Home Sewage Disposal (1924).
Weterdale, J. S. Memorandum on Latrines. Government of Madras,
India (1928).
Savignac, A. L. A new drill used by the Preston Engineering Depart-
ment. Unifruitco. The United Fruit Company, Boston, Massachu-
setts No. 9, 5 (1930) 535.
Jacocks, W. P. A note on a type of latrine suitable for use in Ceylon
villages. Ceylon Journ. Sci. § D., Medical Science, Part 2, 2 (1929).
Yeager, C. H. The bored-hole latrine. Malay. Med. Journ. No. 1, 5 (1929)
1-3.
Yeager, C. H. Practical bored-hole latrine construction. Malay. Med.
Journ. No. 2, 4 (1929) 45-55.
Yeager, C. H. Well pollution and safe sites for bored-hole latrines. Ma-
lay. Med. Journ. No. 4, 4 (1929) 118-125.
Reports of the Rockefeller Foundation.
U. S. Public Health Service Reports.
ILLUSTRATIONS
Plate 1
Reenforced concrete squatting slab designed by J. S. Westerdale for Ma-
dras.
Plate 2
One of the rectangular cement slabs widely used in the Philippine Islands.
The lid and handle shown in AhA is the type formerly used
in Java. The slab is reenforced with £-inch twisted iron.
The cost is less than 2.50 pesos.
Plate 3
Detailed sketch of the elevated water trap.
Plate 4
Detailed sketch of the water trap and slab shown in fig. 46.
Plate 5
Latrine used by the United Fruit Company.
Plate 6
A latrine used in Siam fitted with a sliding lid which must be closed in
order to open the door to get out.
Plate 7
Fig. 1. The rock was shattered covering a radius of ten feet from the
blast holes in our first attempt to install latrines in tuff. Clean-
cut cylindrical latrines are now being blasted daily since the
method was improved.
2. Small cement slab used in Java. Photograph sent by Doctor Hy-
drick who reports favorably on these slabs. They cost less
than a gilder.
TEXT FIGURES
Fig. 1. The Iwan auger, a, Shaft attached to auger arch; 6, a more solid
joint with a nut below the socket. The bolts and nuts are not
necessary if the shaft is welded to the arch.
2. The chain-tong turning handle, made by attaching the handle
a to a Vulcan bijaw chain tong. This is an excellent turning
device and has been used in many places.
264209 14 745
746 The Philippine Journal of Science mi
Fig. 3. Crumbie tongs, one of the most satisfactory inexpensive tongs on
the market, a, Old type; b, improved Crumbie tongs; these
cost a little more, but are worth it.
4. Turning handle designed by Doctor Hamilton. "The two steel re-
tention plates with the grooves on their inner surfaces serve to
prevent the dogs from falling out of place when no pipe is be-
tween them. The plates are bolted together and to the frame of
the body so that the dogs are easily removable and exchanged
when worn or damaged.,, (Drawing from a sketch sent by Doc-
tor Hamilton to the Rockefeller Foundation.)
5. Turning handles, a, The pipe-cross turning handle. This outfit
is usually used in the Philippine Islands for general distribution
"because it is the least expensive satisfactory device we have
tried. The only materials required are two pieces of pipe, a
heavy cross or four-way joint, and a tool-steel pin to transfix
the cross and shaft. The cross slides easily up or down the
shaft; b, a turning handle for use on a shaft drilled with holes
to engage the lugs. This wrench is more expensive than the
pipe-cross handle.
6. Coupling and shaft, a, A solid coupling for shafts made in sec-
tions; two bolts are removed to take the shaft apart; 6, the
bolt shaft, one of the most useful shafts we have used. This
shaft stands more rough use than any other shaft tried. Per-
manent bolts or rivets transfix the shaft at 3-foot intervals.
The turning handles never damage this shaft. The bolt heads
act as lugs for the handles to push against.
7. Turning handles, a, One type of locally made turning handle
for the bolt shaft, hammered out by a blacksmith. These can
also be made of cast iron ; 6, another type of locally made handle
for the bolt shaft. The head of a bolt on the shaft enters
the socket r, and the nut on the opposite end fits into notch
n; c, a turning handle that can be made locally or purchased
ready-made from dealers.
8. A more elaborate turning handle for use on the bolt shaft. This
is an excellent handle but costs 20 pesos to make locally. In
large quantities the cost should be less.
9. The A frame now used instead of a tripod for hoisting augers.
The guy ropes are usually tied to a house, tree, or fence post.
Bamboo is usually used because it is much cheaper in the
Philippines than iron pipe.
10. In some places augers are lifted by direct pull, but the job is too
heavy in most areas. The rope is attached to the arch or low
down on the shaft.
11. Trap doors costing 6 pesos greatly facilitate boring. The auger
shaft is supported by the doors when closed. Stakes can be
driven into the earth at the notched corners to prevent move-
ment of the platform while turning. One of these platform
braces is now included as standard equipment with every auger
used in the Philippines.
46,4 Y eager: Bored-hole Latrines 747
Fig. 12. This tripod is more difficult to transport than the A frames, but
if properly made greatly speeds up boring when several holes
are to be bored within a small area, a, Shaft support. A
knock on the extension lever c releases the auger so that it can
be swung away to be emptied; 6, an iron hook that is a great
time and energy saver. These hooks hold the auger away from
the latrine while being emptied. The hooks are also used on A
frames.
13. An excellent shaft brace designed by Doctor Hamilton. The cost
of production is a disadvantage of this device. (Drawing from
Doctor Hamilton's sketch.)
14. A quadruple expansion brace sold by the Alsdorf Corporation.
The stock size is made to fit the Standard earth auger.
15. A locally made swivel for use on the top of the shaft. We have
used these swivels but prefer unhooking the rope from the shaft
while turning the auger.
16. Devices which can be attached to the auger shaft and adjusted to
any position so that a hook on the hoisting rope can be quickly
hooked on instead of tying and undoing knots; a and b are used
on shafts with holes drilled at intervals such as used with the
pipe-cross handles ; c and d are used on the plain or bolt shafts.
17. Reamer, stone hooks, and grapple. at An undercutting reamer
which is useful in cutting away the sides of latrines below lin-
ings; fortunately it is rarely necessary to use one of these;
6 and c, stone hooks which are useful in removing small bowl-
ders; d, a grapple (redrawn from picture from R. R. Howell &
Co.).
18. Augers suitable for soft mud and silting sand, a, Type made by
many manufacturers of drilling equipment; 6, a heavy dump-
ing auger such as used with machine-driven outfits; this auger
is made by the Gus Pech Co.; c, the Lang auger with sand-bor-
ing screw; this is a good hand auger for boring in sand. See
fig. 31, Howell drop-bottom auger.
19. Sand pumps and bailers which can be purchased from dealers.
These pictures are from the Keystone Drilling Co., Beaver
Falls, Pa.
20. The Iwan post-hole auger fitted with valves for use in silting
sand.
21. A hinge to facilitate turning an auger over so that it can be
dumped.
22. A hinge to facilitate turning an auger over so that it can be
dumped.
23. A hinge for the same purpose as those shown in figs. 21 and 22.
24. An auger designed by A. L. Savignac for the United Fruit Co.
This auger works in soft mud. Note the hinge for dumping.
25. A clay and sand auger that can be made locally. Augers of
similar design are sold by many manufacturers without the flap
valve a. R. R. Howell & Co., Minneapolis, manufacture these
augers. A hinge on the shaft is not needed because the bottom
of the auger swings back on a hinge to empty the contents.
748 The Philippine Journal of Science iski
Fig. 26. A valve auger designed by Doctor Hamilton in Java.
27. A short chisel-bottom bailer designed by the engineering depart-
ment, Sarawak Oil Field, Ltd., Miri, Sarawak. This drill
should be a good one, but is expensive to make and requires
six men to handle effectively.
28. Disc augers. These are probably the cheapest augers made, but
are not as good as other augers mentioned.
29. The Standard auger, sold by the A. J. Alsdorf Corporation.
30. The Lang-London, Ltd., auger. This auger is used in many places
in clay and especially in soft sand. When used in sand a special
screw, shown in fig. 18, c, is attached.
31. Various earth augers manufactured by R. R. Howell & Co. a,
For clay and hard pan; o, for boring and removing core; c and
d, for general boring; e, for loosening and removing stones; /,
for loose sand soil; g, a drop-bottom, fast-cutting auger espe-
cially useful with power-driven machines; hf a spudding jetting
drill used in rock drilling. Blasting is much more rapid for
latrine installation in rock.
32. Animal-driven boring apparatus. Used for many years in well
drilling, but not suitable for rapid latrine boring.
33. The Budda-Hubron machine drill. This is an efficient rotary
drill, and takes less operating space than any machine we have
heard of. It can be used for boring in clay, sand, hard pan,
shale, and frozen ground. The stock machine bores holes 10
feet in depth, but the manufacturers will equip it for boring
20-foot holes. It costs about 2,400 dollars United States cur-
rency.
34. The Gus Pech power-driven machine is a rapid borer, but re-
quires a space 10 by 16 feet for efficient operation. It costs
less than 1,000 dollars equipped for latrine boring.
35. Tools used for blasting latrines in rock, a, Bar used in drilling
blast holes in tuff, adobe rock, and other hard formations; a 3,
drill used with hammer for making blast holes in hard rock;
o, crow bar used for starting latrine or straightening side; c,
long bar used occasionally in deep latrines; d, bamboo bucket
for removing water from blast holes ; e, bamboo brush for clean-
ing mud out of blast holes.
36. Blasting in rock, a to e, Positions of blast holes. In hard rock
five blast holes are drilled at a and b.
37. A hand drill especially designed for making blast holes in tuff,
adobe rock, shale, or other hard formations. It will not cut
hard rock. This drill cuts the holes rapidly and cleans them out
at the same time; it is constantly used now in place of the bar
a, fig. 35, except in the hardest rock where the drill a3 is used.
Manufactured by the Howells Mining Drill Co., Plymouth, Pa.
Cost about 18 dollars.
38. A geared drill for boring blast holes rapidly in very hard forma-
tion except hard rock. Manufactured by Howells Mining Drill
Co., Plymouth, Pa. Cost, 140 dollars.
46,4 Y eager: Bored-hole Latrines 749
Pig. 39. Derrick manufactured by Werf Conrad, Haarlem, Holland. For
rotary, percussion, or free-ball system of boring. This appa-
ratus will handle heavy drills. It is too heavy for general bored-
latrine work.
40. Hand-power spudding and hoisting windlass manufactured by
Werf Conrad, Haarlem, Holland. Believed to be too expensive
(about 200 dollars) and too bulky for general distribution for
latrine boring.
41. Woven-bamboo cylinder partly made. This makes a very satisfac-
tory latrine lining in most places. The section a is a tempo-
rary support to keep the ribs straight while weaving; b is an
enlarged section of the woven-bamboo latrine lining.
42. Various latrine linings, a, A galvanized-iron wire-net lining
which can be used in places where white ants eat the bamboo
or where bamboo is not available; b, stove-pipe double wall,
"wall method" of making a lining. In ordinary soils numerous
holes can be cut into the metal to allow better action. In sil-
ting soils the sections are added one at a time as boring pro-
ceeds. A few dents hold the cylinders together; c, four to six
empty metal oil or tar drums placed end on end are frequently
used as linings for latrines. The heads of the drums are cut
out with a chisel. Wooden or metal strips are nailed to the
drums to make strong joints. These are often used in silting
sand.
43. A steel latrine floor, or squatting plate, designed by Dr. W. P.
Jacocks for use in Ceylon. The cost is about 4 rupees, about
2.60 pesos.
44. Latrine and water closet, a, The bored-hole latrine complete with
superstructure. The lining and cement casing are not used in
soil that does not cave into the latrine. Metal drums are
usually used instead of cement casings where the water rises to
the surface; 6, a cement water closet that can be flushed with
a bucket of water. About two hundred of these water traps
are giving excellent service in the Philippines. Note clean-out
hole in Plate 3. The sloping foot rests make the user sit on
the water closet correctly. The rests throw one off balance if
he attempts to squat backwards. These traps cost about 4 pesos
each. They absolutely eliminate fly and mosquito breeding and
foul odors.
45. Reenforced concrete slab for use with water trap. Elevated type.
46. Latrine equipped with water trap sunk flush with the slab.
This trap costs only 3 pesos. Note the sloping foot rests and
the small vent, which can be opened to relieve pressure in an
air-tight latrine. The washings from the trap fall from the lip
and do not follow the trap down to the wall of the latrine.
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PLATE 7.
THE PHILIPPINE VARIETIES OF ANOPHELES GIGAS
AND ANOPHELES LINDESAYI
By W. V. King
Of the International Health Division, Rockefeller Foundation1
TWO PLATES
Anopheles formosus was described by Miss Ludlow in 1909
from a specimen collected at Camp John Hay, Baguio, Benguet,
on Luzon Island. It was later considered by Christophers to
be a variety of Anopheles gigas Giles. The species has not since
been reported from the Philippines and the larva has not pre-
viously been described. The following description of this stage
is based on an examination of 35 larvse collected in the same
locality in April and May, 1929, and in May, 1931. One of the
three collections, that of May, 1929, was made for me by Ser-
geant J. F. Rhodes, of the United States Army Medical Corps.
The larvse were taken at an altitude of about 4,700 feet.
ANOPHELES GIGAS var. FORMOSUS Ludlow, 1909.
Larva. — Inner anterior clypeal hairs (Plate 1, fig. 1) usually
simple but occasionally split into two, the bases close together ;
the outer clypeal hairs one-half or more the length of the inner
and branched two to six times, sometimes simple; posterior
clypeal hairs nearly as long as the outer and somewhat closer
together, branched toward the base from three to eight times,
the usual number four or three. Occipital hairs large, the in-
ner branched from four to twelve times, usually from eight to
ten ; the outer branched from seven to sixteen times with counts
of nine, ten, or eleven the most frequent. Inner anterior sub-
median thoracic hair with three to ten branches ; middle anterior
hair much longer with from eight to fifteen branches (Plate 1,
fig. 3). Palmate hairs lacking on the thorax and first two
abdominal segments, being represented by ordinary branched
hairs, the one on the thorax with four to ten branches (Plate
"In cooperation with the Bureau of Science and the Philippine Health
Service.
751
752 The Philippine Journal of Science imi
1, fig. 5). Well-developed palmate hairs present on abdominal
segments III to VII, the individual leaflets more or less bluntly
tapered, without filaments, and either smooth or with a few ser-
rations on the edges (Plate 1, fig. 7). All of the antipalmate
hairs (hair 2 of Martini, 1923) multiple, those on segments
III to VII having from three to eight branches with five the
most frequent number; antipalmate hair on segment II with
from four to eleven branches, seven and eight being the most
frequent. The long lateral hairs on abdominal segments IV
and V usually 3-branched but vary from two to five in the
series; this hair lacking on segment VI and is represented by
a very short, branched tuft. Pecten (one specimen) with six
long and fifteen short teeth.
The larvae were collected at the grassy margins of pools in
stream beds and on one occasion along the edge of a large rock
in a well-shaded stream pool.
Adult females reared from Baguio larvae agree in general
with Ludlow's original description except that the fringe spot on
the posterior margin of the wing occurs between veins 5.2 and
6 instead of between the forks of the 5th vein as stated. The
type specimen now in the United States National Museum in
Washington has it in this position also so that the original de-
scription was in error.
The palpi have three very narrow white bands and the apex
is more or less pale, some specimens having distinct yellowish
scales, others only pale apical hairs. The 6th vein of the wing
has a white scaled area two or three times the length of the
apical dark spot, the subapical costal white spot is absent and
the extreme base of the costa is white (Plate 2, fig. 1) .
Slight differences in adult markings have been given for the
several varieties of this species, and Christophers (1931) has
recently published a revised summary of the group. The varie-
ties recognized, are:
Anopheles gigas Giles, 1901, type form from southern India.
Var. formosus Ludlow, 1909, from Luzon, Philippine Islands (not For-
mosa as listed by Christophers).
Var. simlensis James, 1911, from the Western Himalayas.
Var. refutans Alcock, 1913, from Ceylon.
Var. baileyi Edwards, 1929, from Western China, Eastern Himalayas,
Assam, Burma and Tibet.
(Anopheles edwardsi Yamada, 1924, from Japan, is considered a dis-
tinct species.)
The Philippine form appears, from palpal and wing markings,
to resemble variety refutans more closely than any of the others.
461 King; Varieties of Anopheles 753
Both have pale-tipped palpi and the wings are without fringe
spots except in the area between veins 5.2 and 6. The two forms
probably differ in the scaling of the extreme base of the costa,
which Christophers shows to be dark scaled in variety refutans.
He also records the occurrence of this character in variety
formosus, but all Philippine specimens examined by me have a
white scaled area at the extreme base of the costa, nearly equal
to or longer than the succeeding (inner accessory) dark spot, as
shown in the accompanying illustration.
The larval characters of the Ceylon form so far as given by
Carter (1925) also appear to be similar to those of the Philippine
variety except that the post-clypeal hairs are said to be simple
or with two or three divisions, whereas none of the specimens
examined in this series of var. formosus have hairs with less
than three branches.
Male genitalia of var. formosus (Plate 2, figs. 3 and 5). —
Inner parabasal spine of side piece broad and flattened for
entire length, about 0.11 mm long; outer spine 0.14 mm
long, more slender and tapered to a long point. Outer lobe
of harpagones (claspette) with five to seven unfused spatulate
filaments or blades, the internal one somewhat longer than the
others, arising from a separate prominence. Length of latter
0.11 mm and the longest one of the others 0.09 mm. The indi-
vidual blades are bluntly rounded and end in a minute thornlike
point. On the inner lobe of the harpagones a group of three
hairs placed close together, the outer 0.09 mm long, the middle
one very short and the inner one, at the apex of the lobe, 0.14
mm long. In one of three specimens examined, the middle hair
on one side is also long. Mesosome (theca) with five to seven
leaflets, the longest one about 0.05 mm, the others progressively
shorter. Under high magnification most of the leaflets show
serrations. The ventral processes of the 9th segment are not
apparent except possibly as small humps.
The genitalia of the Philippine variety appear to differ slightly
from that of A. gigas as described by Christophers (1915) in
the hairs of the harpagones and in the ventral processes of the
9th segment.
ANOPHELES LINDESAYI var. BENGUETENSIS var. nov.
A collection made for me by Sergeant Rhodes at Camp John
Hay in May, 1929, contained, in addition to larvae of A. maculatus
and A. gigas var. formosus, several specimens of this species,
which is the first record of its occurrence in the Philippines.
Sergeant Rhodes said that the larvae were collected along the
754 The Philippine Journal of Science iwi
edge of a well-shaded stream among leaves and debris or at the
side of rocks. Additional collections in the same locality have
been made by Mr. F. E. Baisas during 1930 and by myself in
May, 1931.
Adults of A. lindesayi have unhanded palpi and tarsi and are
readily identified by the presence of a broad white band on the
distal half of the hind femora. The wing markings of the
female of the Philippine variety are shown in the accompanying
illustration (Plate 2, fig. 2). The white-scaled spots at the
tips of veins 4,2, 5.2, and 6 and the fringe spot at 5.2 are evident-
ly constant, being present in each of the seventeen specimens
examined. No white scales occur at the ends of veins 2.2, 3,
4.1, or 5.1. The wing fringe opposite veins 6 and 4.2 is variable
and may be either dark or slightly pale.
The markings of the wing are therefore slightly different
from any of the varieties of this species as listed by Christophers
(1931). The forms recognized by him are:
A. Imdesayi* Giles, 1900, type form, from the Himalayas,
Var. japonims Yamada, 1918, from Japan.
Var. pleccau Koidzumi, 1920, from Formosa (provisionally retained).
Var. nilgiHcu8 Christophers, 1924, from South India.
Var. cameronensis Edwards, 1929, from the Federated Malay States.
The Philippine form probably comes closest to var. japonicus.
The scaling at the termination of the veins in the latter variety
is, however, considerably more variable. Yamada (1924) notes
the occurrence of white spots at the ends of veins 3, 4.2, 5.1, 5.2,
and 6 and adds that those at 3, 4.2, and 5.1 may or may not
appear according to the specimens. Fringe spots also may or
may not occur opposite veins 4.2, 5.1, and 5.2.
The fore and mid femora of the Philippine specimens have
distinct white bands or rings at the base, equal to or less than
the diameter of the femora in extent. The bands on the hind
femora are variable but usually wider. In a number of speci-
mens the ventral white is two to three times the diameter of
the femoral joint and the dorsal white somewhat shortened (one
to two times the diameter.) In certain specimens the white is
practically the same above and below, while in two specimens
the black scaling extends nearly to the base above and the white
scaling ventrally is more extensive — between one-fifth and one-
sixth of the length of the femora by measurement in one speci-
a Named after Captain Lindesay but originally spelled lindesaii. The
changed spelling followed here was made, I believe, by Blanchard in 1905.
46,4 King: Varieties of Anopheles 755
men. The arrangement of white seems to be more of less similar
to that of var. earner onensis, but, according to Christophers, that
form has none of the wing veins from 2.2 to 5.2 white-tipped.
Male genitalia. — Inner parabasal spines stout, flattened and
recurved, 0.10 mm long (one measurement) ; outer stout and
somewhat flattened toward end, 0.14 mm long. Outer lobe of
harpagones with three bladelike filaments subequal in length and
broadest near tip, the longest about 0.07 mm; inner lobe with
a stout hair at apex, 0.10 mm long, and well separated from
this internally a slenderer hair, 0.07 mm long (Plate 2, fig. 6) .
Mesosome with very many slender leaflets (Plate 2, fig. 4),
eighteen or nineteen on each side in three specimens, counted
after separating the mesosome from the hypopygium and flat-
tening under a cover glass. Some of the leaflets are serrated
along the side and some are split at the tip.
The harpagones of A. lindesayi as described by Christophers
(1915) have two bladelike spines on the outer lobes, instead
of three, and a much smaller number of leaflets on the meso-
some— about five as compared to eighteen or so in var. benguet-
ensis.
Larva. — The characters recorded for one of the larvae from
the original collection are as follows : Inner anterior clypeal hairs
long, simple and close together ; outer clypeals simple, less than
half the length of the inner ; postclypeals about as long and as
widely separated as the outer, two branches on one side and
three on the other (Plate 1, fig. 2) . Inner occipital hairs simple
on one side, forked on the other ; outer occipitals three and four
branched. Inner anterior submedian thoracic hairs (Plate 1,
fig. 4) with thirteen and fourteen branches; middle hairs longer,
with fourteen and twenty branches. Thoracic palmate tuft de-
veloped, the leaflets being slenderer than those on the abdomen
(Plate 1, fig. 6) ; abdominal palmate tufts developed on segments
II to VII, the leaflets with filaments (Plate 1, fig. 8) . The lateral
hairs on abdominal segment IV, three branched, on segment V,
three and two branched and absent on segment VI.
From other specimens collected and examined by Mr. Baisas,
the postclypeal hairs are sometimes simple, although two or three
divisions is the usual number. The antipalmate hairs on seg-
ment II have about six branches; on segment III usually five or
six; on segments IV and V, single; on segment VI usually three
and on segment VII, five.
The larvae are pigmented and mottled on the dorsal surface of
the thorax and abdomen, and wide pigmented bands completely
756 The Philippine Journal of Science
surround abdominal segments II, IV, VI, VII, and VIII, giving
the larvse a characteristically striped appearance and distinguish-
ing them from other species with which they occur. The band-
ing still appears after preservation in formalin.
Type female reared from larva collected at Baguio, Benguet
Subprovince, Luzon, Philippine Islands, May 20, 1931. Taken
at an altitude of about 4,700 feet.
REFERENCES
Alcock, A. Synopsis of the anopheline mosquitoes of Africa and of the
Oriental region. Journ. Lond. Sch. Trop. Med. 2 (1913) 153-166.
Carter, H. F. The anopheline mosquitoes of Ceylon. Part I. The dif-
ferential characters of the adults and larvae. Ceylon Journ. Sci. §
D, Med. Sci. 1 pt. 2 (1925).
Christophers, S. R. The male genitalia of Anopheles. Ind. Journ. Med.
Res. 3 (1915) 371-394.
Christophers, S. R. Some Himalayan and Peninsular varieties of Indian
species of Anopheles. Ind. Journ. Med. Res. 12 (1924) 11-13.
Christophers, S. R. Studies on the anopheline fauna of India. (Parts
I-IV). Reeds. Mai. Surv. Ind. 2, No. 2 (1931) 305-332.
Giles, G. M. Handbook of the Gnats or Mosquitoes. 1st. ed. (1900).
Giles, G. M. Descriptions of four new species of Anopheles from India.
Ent. Month. Mag. 2d ser. 12 (1901) 190-198.
James, S. P„ in James and Liston. A monograph of the Anopheline mos-
quitoes of India, 2d ed. (1911).
Koibzumi, M. Daiwan Kenkyujo Hokoku 8 (1920) 17, 28, and 34. (Cited
by Yamada, 1924.)
Ludlow, C. S. Mosquito comment. Canad. Ent. 41 (1909) 22.
Martini, E. Uber einige fur des System bedentungsvolle Merkmale der
Stechmxiehen. Zool. Jahr., Abt. fur Syst. Geo. Biol, der Tiere, 46
(1923) 517-590.
Yamada, S. Eiseigaku Densenbyogaku Zasshi 13 (1918) 689. (Cited by
Yamada, 1924.)
Yamada, S. A revision of the adult anopheline mosquitoes of Japan. Sci.
Kept. Govt. Inst. Inf. Dis. 3 (1924) 215-241.
ILLUSTRATIONS
[The illustrations are from camera lucida drawings made by F. E. Baisas and F. del Rosario.]
Plate 1
Fig, 1. Clypeal hairs of A. gigas var. formosus.
2. Clypeal hairs of A. lindesayi var. benguetensis.
3. Anterior submedian thoracic hairs, left side, of var. formosus.
4. Anterior submedian thoracic hairs, right side, of var. benguetensis.
5. Branched hair in place of thoracic palmate of var. formosus.
6. Thoracic palmate of var. benguetensis.
7. Two leaflets from an abdominal palmate tuft, segment IV of var.
formosus.
8. Two leaflets from an abdominal palmate tuft, segment IV of var.
benguetensis.
Plate 2
Fig. 1. Wing of var. formosus.
2. Wing of var. benguetensis.
3. The leaflets from one side of the mesosome of the male genitalia
of var. formosus.
4. The leaflets from one side of the mesosome of var. benguetensis.
5. Half of the harpagones of var. formosus.
6. Half of the harpagones of var. benguetensis.
757
King: Varieties op Anopheles.]
[Philip. Journ. Scl, 46, No. 4.
PLATE 1.
King: Varieties op Anopheles.]
[Philip. Journ. Scl, 46, No. 4.
PLATE 2.
THE USE OF THE ANTENNJE AS A MEANS OF DETER-
MINING THE SEXES IN LEUCOPHOLIS IRRORATA
ADULTS (COLEOPTERA, SCARABJEIDJE)1
By A. W. Lopez
Chief Entomologist, Research Bureau, Philippine Sugar Association
ONE PLATE
INTRODUCTION
The flying season of Leucopholis irrorata beetles in Occiden-
tal Negros normally occurs during the latter part of April, all
of May, and the first part of June.
During the season of 1930, this entomology department ex-
amined 1,663 beetles in order to determine the sex ratio and the
egg content of beetles collected in the field, for the purpose of
securing data on the soundness of the collecting campaign prin-
ciple. Because neither the writer nor other local entomologists
could differentiate between the sexes at sight, it was necessary
to dissect every individual in order to determine its sex, with a
consequent expenditure of a great amount of time.
At the start of the 1931 season it became apparent to the
writer that males could be separated from females through cer-
tain characteristics inherent in the antennse. It is now possible
to take a group of beetles on which data are desired, and rapidly
and accurately to pick out the males and females. The former
may then be counted, and only the females dissected. This dif-
ferentiation cannot be practiced immediately, but a small
amount of practice will enable one to so differentiate rapidly.
MATERIAL AND METHODS
One antenna and the elytron from the same side were removed
from a live L. irrorata beetle, and the club of the former and the
total length of the latter were measured with a stage micrometer
to one-tenth of a millimeter. An assistant then dissected the
*The most important sugar-cane white grub (buc-an) of the Philip-
pines. ?59
760
The Philippine Journal of Science
1931
beetle positively to determine its sex. One hundred antennal
clubs and elytra were so measured for each sex.
In making the drawings of the extended antennal clubs, some
difficulty was encountered because they immediately became com-
pact when the antennae were severed. Submersion in 80 per
cent alcohol or concentrated acetic acid for a short time caused
them to extend themselves in approximately the normal manner.
The drawings were made with the aid of a camera lucida.
THE DIFFERENTIATION OF THE SEXES
The length of the male antennal club (Plate 1, fig. 2) as
deduced from the measurement of one hundred individuals,
averages 1.87 millimeters ± 0.0076 millimeter 2 and the female
antennal club Plate 1, fig. 1) averages 1.3 millimeters ± 0.0073
millimeter, the male antennal club being approximately 0.57 mil-
limeter longer than that of the female. While the difference is
not great it is readily perceptible to the naked eye.
The elytra were measured in order to ascertain whether or
not the size of the beetle had an appreciable influence on the
length of the antennal club. If the size of the beetle should
influence the size of the club, then a female that happened to be
larger than a male would have a longer club and the differen-
tiation could not be made. A summary of the results of the
measurement of the clubs and of the elytra, which are considered
indicators of the size of the beetle, is shown in Table 1.
Table 1. — Showing summary of results of club and elytra measurements
in Leucopholis irrorata.
Average length in 100
individuals.
Sizes.
Elytron.
Antennal club.
Shortest
elytron.
Antennal
club.
Longest
elytron.
Antennal
club.
Male
Female
mm.
18.3±0.0494
18.9±0.0615
mm.
1.87±0.0076
1.3±0.0078
mm.
16.5
16.4
mm.
1.8
1.2
mm.
19.6
21.0
mm.
1.9
1.4
In Table 1, the female elytron is shown to be approximately
0.6 millimeter longer than that of the male, while the antennal
club is approximately 0.57 millimeter shorter than that of the
male, as mentioned above. It may also be noted in the table
that the male beetle with the shortest elytron had a club about
* Probable error for the mean.
46,4 Lopez: Antennse in Leucopholis 761
0.07 millimeter shorter than the average, and that the male with
the longest elytron had a club about 0.1 millimeter longer than
the average. In the females there is only a difference of about
0.1 millimeter in the length of the antennal club either way from
the average in the beetle with the shortest and in the one with
the longest elytron. It is thus seen that the size of the beetle
apparently does not influence the length of the antennal club
to any appreciable extent.
In addition to making the differentiation between the sexes
by the difference in length of the antennal clubs alone, the dif-
ference in the contour between the posterior edges of the male
and female extended antennal clubs can also be made use of.
In the female extended antennal club (Plate 1, fig. 3) at point
a9 the posterior edges of the first two lamellae form nearly a
smooth curve with the last funicular segment (Plate 1, fig. 1,
/), while in the male (Plate 1, fig. 4) at point a, the posterior
edges of the first two lamellae do not form a smooth curve with
the last funicular segment, but a distinct drop is noticed, giving
the male extended club a characteristic appearance different from
that of the female extended club.
The males and females of Lepidiota pruinosa may be separated
in a manner identical with the above.
264209 15
ILLUSTRATION
[a, Reference point ; c, club ; /, funicle ; I, lamellie ; p, pedicel ; «, scape.]
Plate 1
Pig. 1. Right antenna of female Leucopholis irrorata, X 30.
2. Right antenna of male L, irrorata, X 30.
3. Extended right antennal club of female L. irrorata, X 30.
4. Extended right antennal club of male L. irrorata, X 30.
763
Lopez: Antennae in Leucopholis.]
[Philip. Journ. Scl, 46, No. 4.
PLATE 1.
INDEX
[New names and new combinations are printed in boldface.]
Aberia gardneri, 516.
Abies magnifica, 4.
sibirica, 4.
Acanthocephala Rudolphi, 538, 582, 584.
Acanthostigma bambusae v. Hoehn., 501.
Acer niveum, 515.
sp., 515.
Acerbia maydis Rehm, 501.
Achrionota Pasc, 425, 435, 442.
bilineata Pasc, 389, 425, 427, 442.
spinifer Kin., 389, 425, 427, 442.
Aerostichum zollingeri Kze., 220.
Actiniptychus undulatus (Bail.) Ralfs, 97.
Actinodothis piperis Syd., 485.
Adelphomyia apoana Alex., 457, 458.
carbonicolor Alex., 458.
nebulosa (de Meij.), 459.
paucisetosa Alex., 458, 459.
Aecidium, 518.
alchorneae Sacc, 482.
banosense Syd., 482.
blumeae P. Henn., 482.
clerodendri P. Henn., 482.
elaeagni-latifoliae Petch., 482.
flavidum Berk. & Br., 482.
kaernbachii P. Henn., 482.
lagunense Syd., 482.
luzoniense P. Henn., 482.
nummulare Berk., 482.
paederiae Diet., 483.
rhytismoideum Berk. & Br., 483.
uvariae-rufae P. Henn., 483.
Aedes, 43, 595, 604-606.
jegypti Linn., 40, 595, 602, 607.
Aegiceras corniculatum, 527.
Afzelia bijuga, 498.
Agriorrhynchus Power, 418, 435, 441.
ignarius Kin., 388, 418, 422, 441.
Aithaloderma clavatisporum Syd., 487.
Aids spinosa, 555.
Albizzia lebbek, 528.
procera, 481.
Alchornea javanica, 528.
rugosa, 482, 503, 528.
Aldona stella nigra Rac, 515.
ALEXANDER, CHARLES P., New or little-
known Tipulidae from the Philippines
(Diptera), X, 9; XI, 269; XII, 447.
Allaeodromus, 892.
Allium cepa-, 531.
sativum, 531.
Allophyllum dimorphum, 535.
Alocasia indica, 504.
Alsophila, 515.
Alstonia, 485.
macrophylla, 489.
scholaris, 485, 528.
Alternaria brassicae (Berk.) Sacc, 533.
Amorphocephalini, 390, 415, 431, 433, 434,
440.
Amorphophallus campanulatus, 480.
Amphicordus K. M. Heller, 422, 434, 441.
improportionalis Heller, 389, 422, 441.
Amphisphaeria arengae Rehm, 503.
schizostachyi Rehm, 503.
Amphisphaeriaceae, 503.
Amphoromorpha entomophila Thaxter, 518.
Anagyrus sp., 222.
Ananas comosus, 489.
sativa, 489.
sativas, 489.
sativus, 489.
Andropogon aciculatus, 484.
eitratus, 480.
halepensis, 480, 500.
halepensis var. propinquus, 500.
sorghum, 480, 484, 488, 500, 507, 534.
Anepsiotes Kin., 424, 435, 442.
luzonicus Calabr., 389, 424, 442.
nitidicollis Calabr., 389, 424, 442.
Anisolabis annulipes, 555.
Anisopia farinalis, 555.
Anoa, 142.
mindorensis, 142.
Anona muricata, 525.
Anopheles, 43, 639, 640, 645, 647.
aconitus var. filipinaa Manalang, 40.
albimanus, 657.
bancrofti var. pseudobarbirostris (Lud-
low), 646.
barbirostris (van der Wulp), 646.
culicifacies, 247, 641.
edwardsi Yamada, 752.
formosus, 751.
fuliginosus (Giles) 646.
funestus Giles, 47, 49, 56-58, 249, 253,
363, 365-367, 372-375, 644.
gigas Giles, 751, 752.
gigas Giles var. baileyi Edw., 752.
765
766
Index
Anopheles — Continued.
gigas Giles var. formosus Ludlow, 751-
753.
gigas Giles var. refutans Alcock, 752,
753.
gigas Giles var. simlensis James, 752.
hyrcanus var, sinensis (Wied.), 646.
kochi (Donitz), 645, 646.
lindesayi, 751, 754, 755.
lindesayi var. benguetensis King, 753,
765.
lindesayi var. cameronensis Edw., 754,
755.
lindesayi var. japonicus Yamada, 754.
lindesayi var. nilgiricus Christophers,
754.
lindesayi var. pleccau Koidzumi, 754.
ludlowi Theo., 40, 374, 642.
maculatus Theo., 40, 642, 753.
maculipennis Mg„ 53, 363, 366,
minimus (Theo.), 644, 646.
philippinensis (Ludlow), 374, 646.
punctipennis, 364.
quadrimaculatus Say, 344, 364-367.
subpictus, 645.
tessellatus (Theo.), 645, 646.
vagus Donitz, 40, 874, 645, 646.
Antennae, use of, as a means of determining
the sexes in Leucopholis irrorata adults
(Coleoptera, Scarabaeidae ) , 759.
Anthostoma eumorphum, 505.
Anthostomella arecae Rehm, 505.
arengae (Rac.) Rehm, 505.
atronitens Rehm, 505.
calami Rehm, 505.
calocarpa Syd., 505.
eocoina Syd., 505.
coryphae Rehm, 505.
coryphae Rehm f. minutissima Rehm,
505.
discophora, 506.
donacina Rehm f. arengae Rehm, 505.
eumorpha (Sacc, & Paoli) Rehm, 505.
grandispora Penz. & Sacc, 506.
lucens Sacc, 506.
micraspis (Berk.) Sacc. & Trav., 506.
mindorensis, 505.
mirabilis (B. & Br.) v. Hoehn., 506.
pandani (Rehm) Syd., 506.
uberiformis Rehm, 506.
Anthraxonis quartiniani, 483.
Antidesma bunius, 487.
Apherodothis arengae (Rac.) Shear, 501.
Aphodius, 578.
Aphysa desmodii Syd., 523.
Apiospora apiospora (Dur. & Mtg.) v.
Hoehn., 496, 497.
camptospora Penz. & Sacc, 496.
carbonacea Rehm, 497.
luzonensis P. Henn., 496, 497.
montagnei, 496.
Apiphragma ( Polyphragma ) nigrotibiata,
463.
Apium jrraveolens, 580.
Apocemus Calabr., 423, 435, 441.
ignobilis Kin., 389, 423, 424, 441.
Apterorrhinus Senna, 429, 435, 442.
albatus Kin., 389, 429, 432, 442.
compressitarsis Senna 389, 429, 432, 442.
Arachis hypogaea, 529, 532.
Araiorrhynchus, 414.
Arcyria carnea G. Lister, 86, 92.
cinerea Pers., 85, 92.
denudata Wettstein, 85, 91.
Ardisia, 487.
Areca catechu, 505, 510, 511, 524, 527-529,
535.
Arenga, 503, 505, 517.
ambong, 536.
mindorensis, 502.
saccharifera, 490, 501, 510, 522, 523.
Arrhenodini, 390, 418, 422, 432-434, 441.
Artemisia sp., 88.
Arthraxonis sp., 483.
Artocarpus, 495, 519.
communis, 495, 520, 521, 531.
incisa, 520, 521, 531.
integra, 520, 521, 534.
integrifolia, 520, 521, 534.
Artotrogus, 518.
Asaphepterum Kin., 397, 433, 437.
formosanum Kin., 386, 397, 400, 437.
Ascari3, 564, 578.
Aschersonia, 493.
cinnabarina P. Henn., 526, 527.
confluens Henn., 493, 526.
javanica, 527.
lecanioides P. Henn., 527.
napoleonae, 527.
novo-guineensis, 527.
paraensis Henn., 527.
phthurioides, 526.
pisiformis, 527.
placenta B. & Br., 527.
samoensis Henn., 493, 527.
sclerotoides Henn., 493, 527.
Ascospora vanillae Rehm, 503.
Aspergillus delacrioixi Sacc. & Syd., 530.
flavus Link., 530.
periconioides Sacc, 530.
sp., 221.
Aspidium, 491.
Asplenium bulbiferum, 215.
bullatum, 215.
finlaysonianum Hook., 215.
finlay8onianum Wall., 214, 215.
fiaccidum, 215.
hookerianum Wall., 214.
integerrimum Hook. & Grev., 214.
macrophyllum Sw„ 214.
tripinnatifidum Copel., 215.
Asterina, 490.
breyniae Syd., 488.
breyniae Yates, 488.
capparidis Syd. & Bult., 488.
cassiae Syd., 488.
colliculosa Speg., 488.
decipiens Syd., 488.
Index
767
Asterina — Continued.
dilleniae S*yd., 488.
elmeri Syd., 488.
gmelinae Sacc, 488.
lawsoniae P. Henn. & Nym., 480.
laxiuscula Syd., 489.
lobata Syd., 489.
opposita Syd., 489.
pandani, 490.
pipturi Syd., 489.
pusilla Syd., 489.
sponiae Rac, 489.
Asterinella calami Syd., 489.
luzonensis Syd., 489.
obesa Syd., 489.
stuhlmanni (Henn.) Theiss., 489.
Asterionella frauenfeldii Gran., 111.
glacialis Castr., 111.
japonica Cleve, 80, 111.
pynedraeformis Grev., 111.
Asteroma phaseoli Brun., 520.
Astilbe philippinensis, 480.
Astrociptis mirabilis B. & Br., 506.
Astronia, 526.
Astrosphariella fusispora Syd., 506.
Ateuchus, 578.
Athyrium blumei, 214.
costulisorum, 214.
ophiodontam Copel., 214.
tenuifolium, 214.
umbrosum, 214.
Atopomorphus Kin., 398, 433, 487.
schultzei Kin., 386, 398, 400, 487.
Aoerswaldia, 502, 505, 506.
decipiens, 505.
examinans (Mont. & Berk.) Sacc, 494.
gigantochloae Rehm, 494.
Aulacostroma palawanense Syd., 494.
Automobiles, decay of wood of, in the Tro-
pics, 189.
Averrhoa carambola, 581.
Avian malaria studies, I, 805; II, 347; III,
651.
Bacillus coli. 611, 613, 616, 619, 736, 787,
739.
fluorescens, 735.
prodigiosus, 738.
proteus, 735.
subtilis, 613, 619.
Backet, see buakat.
Bacteriastrum, 95.
comosum Pavil. var. hispida (Castr.)
Ikari, 109.
hyalinum Lauder, 108, 109.
minus Karst., 109.
varians Lauder, 108.
varians Lauder var. hispida (Castr.)
Schroder, 109.
varians var. hyalina Lauder, 108.
wallichi Ralfs var. hispida Castr., 109.
Badhamia mandshurica Skv., 85, 86.
BAKER, C. F., Second supplement to the
list of the lower fungi of the Phil-
ippine Islands, 479.
Bakerophoma sacchari Diedicke, 520.
Balansia claviceps Speg., 494.
Balladyna velutina (Berk. & Curt.) ▼.
Hoehn., 485.
Bambusa, 493, 496, 497, 504, 506-608, 610,
512, 516, 522.
blumeana, 492, 499, 500-503, 506, 507,
510, 513, 515.
longinodis, 533.
vulgaris, 496, 503, 507, 509, 516.
sp., 496, 510, 513, 583.
Barleria cristata, 511.
Barya salaccensis, 493.
Baryrrhynchus Lac, 419, 435, 441.
schroederi Kin., 388, 419, 420, 422, 441.
Bauhinia, 513, 514.
comingiana, 491.
malabarica, 531.
Beetles, 555, 583.
dung, 578.
Belopherini, 390, 422, 424, 432, 433, 435,
441.
Beta vulgaris, 531.
Biddulphia aurita Brebisson var. oriental!!
Mereschkowsky, 109.
biddulphiana (Sm.) Boyer, 110.
longicornis Grev., 110.
pulchella Gray., 110.
sinensis Grev., 109.
Blaps mucronata, 588.
Blatta orientalis, 556, 578.
Blumea balsamifera, 482.
Botryodiplodia anceps Sacc. & Syd., 520.
curta Sacc, 520.
Botryosphaeria minuscula Sacc, 511.
Brassica chinensis, 88, 89, 531.
culta, 533.
pekinensis, 531.
sinensis, 531.
spp., 581.
Brenthini, 483.
Brentus bisulcatus F., 406.
Breynia cernua, 488.
Briar dia maquilingiana Rehm, 517.
Bridelia glabrifolia, 481.
stipularis, 504.
Broomella zeae Rehm, 491.
Buakat, 257.
Bubalus indicus, 142.
mindorensis, 142.
Bulgariacese, 515.
Bulgariastrum caespitosum Syd., 515.
Csenorychodes Kin., 421, 435, 441.
serrirostris F., 389, 421, 422, 441.
splendens Kbm., 389, 421, 422, 441.
Calamus, 490, 503, 505, 507, 508, 511. 518.
sp., 489.
Calanthes, 528.
768
Index
Callicarpa cana, 485.
sp., 485.
Calodromini, 390, 392, 481-438, 486.
Calodromus Guer., 892, 432, 436.
crinitus Kin., 386, 392, 400, 436.
mellyi Guer., 386, 392, 400, 486.
Calonectria copelandii P. Henn., 491.
hibiscola P. Henn., 492.
meliae A. Zimm., 491, 492.
sulcata Starb., 491.
Calopeziza mirabilis Syd., 515.
Camelia sativa, 518.
Campium subsimplex (Fee) Copel., 220.
Canarium villosum, 498.
sp., 489, 498, 515.
Canavalia, 528.
ensiformis, 517.
gladiata, 508, 517, 528, 581.
Capillariinse Railliet, 564.
Capnodiaceae, 487.
Capnodium footii Berk. & Desm., 487.
Capparis horrida, 488, 504, 508, 523, 624,
530, 535.
irosinensis, 488.
micracantha, 488, 533.
sepiaria, 515.
Capsicum annuum, 523, 526, 528, 586.
Carcara erosa, 482.
Carex sp., 88.
Carica papaya, 504, 507, 518, 519, 521, 522,
528, 530, 534.
Carissa arduina, 529.
Carthamus tinctorium, 533.
Garyota rumphiana var. philippinensis, 501.
sp., 504.
Cassava, 627.
Cassia occidentale, 532.
Catacauma apoense (Syd.) Theiss. & Syd.,
497.
aspideum (Berk.) Theiss. & Syd. f.
ficifulvae (Koord.) Theiss. & Syd., 497.
aspideum (Berk.) Theiss. & Syd. f. spi-
nifera (Karst. & Har.) Theiss. & Syd.,
497.
circinatum (Syd.) Theiss. & Syd., 497.
elmeri (Syd.) Theiss. & Syd., 497.
euryae (Rac.) Theiss. & Syd., 497.
garciae Theiss. & Syd., 497.
infectorium (Cke.) Theiss. & Syd., 498.
kaernbachii (P. Henn.) Theiss. & Syd.,
498.
lagunense (Syd.) Theiss. & Syd., 498.
pterocarpi (Syd.) Theiss. & Syd., 498.
sanguineum (Rehm) Theiss. & Syd., 496.
valsiforme (Rehm) Theiss. & Syd., 498.
Cediocera Pasc, 425, 485, 442.
tristis Senna, 389, 425, 427, 442.
Cenangiacese, 515.
Cenangium blumeanum Rehm, 515.
Centeter, 129.
cinerea, 129.
Centotheca latifolia, 493, 494.
Ceratiomyxa fruticulosa, 86.
fruticulosa Macbr. var. flexuosa Lister,
86.
Ceratophyllus fasciatus, 556.
Cercospora acerosum Dickh. & Hein., 580.
apii Fres., 530.
armoraciae Sacc, 531.
artocarpi Syd., 581.
bauhiniae Syd., 531.
beticola Sacc, 531.
brassicola Henn., 531.
canavaliae Syd., 531.
coffeicola Berk. & Cke., 531.
cruenta Sacc, 531.
duddiae Welles, 531.
gliricidiae Syd., 531.
henningsii Allesch., 531.
lactucae Stevenson, 531.
lactucae Welles, 531.
litseae-glutinosae Syd., 531.
lussoniense Sacc, 532.
mangiferae Koord., 532.
manihotis P. Henn., 532.
melongenae Welles, 532.
nicotianae Ell. & Evht., 532.
occidentals Cke. var. cassiocarpa Sacc,
532.
overrhoi Welles, 531.
pachyderma Syd., 532.
pahudiae Syd., 532.
pantoleuca Syd., 532.
personata (B. & C.) Ell., 532.
puerariae Syd., 532.
sesami A. Zimm., 532.
stizolobii Syd., 532.
subsessilis Syd., 533.
tiglii Henn., 533.
ubi Rac, 533.
Cercosporina carthami Syd., 533.
Cerobates Schoenh., 402, 434, 437.
adustus Senna, 387, 402, 405, 438.
jequalis Kin., 387, 402, 405, 438.
angustipennis Stenna, 387, 402, 403, 405,
438.
clinatus Kin., 387, 403, 405, 437.
costatus Kin., 387, 403, 405, 438.
formosanus von Schonf., 387, 403, 405,
438.
grouvellei Senna, 387, 403, 405, 488.
sexsulcatus Motsch., 387, 402-405, 438.
sumatranus Senna, 387, 404, 405, 438.
tristriatus F., 387, 402, 404, 405, 438.
Ceropegia sp., 482.
Cestoda Rudolphi, 546.
(s. str.) Monticelli, 546.
Cestodes, 538, 584.
Cestodiscus sol Grun., 97.
Ceuthocarpon depokense Penz. & Sacc, 506.
punctiforme Sacc, 506.
talaumae Rehm, 506.
Chaetoceras, 95.
affine Lauder, 106, 107.
angulatum Schtitt, 107.
Index
769
Chaetoceras — Continued .
atlanticum Cleve, 105.
atlanticum var. tumescens Grun., 105.
boreale Bail., 101.
boreale Cleve, 101.
boreale var. brightwellii Cleve, 101.
cellulosum Lauder, 100.
commutatum Cleve, 79.
compactum Schtitt, 105.
compressum Lauder, 78, 104.
compressum var. gracilis Hustedt, 78.
constrictum Gran., 79.
contortum Schutt, 104.
criophilum Castr. forma volans (Schutt)
Gran, 78, 106.
dadayi Pavil., 105.
decipiens Cleve, 79, 104.
didymum, 103.
didymum Ehrenb. var. anglica Gran, 78,
103.
didymum Ehrenb. var. genuina Gran,
103.
didymum var. longicruris Cleve, 108.
dispar Castr., 105.
distans Cleve, 79.
distans Ostenf., 79.
distans var. laciniosa Schutt, 80.
distichum Schutt, 106.
furca Cleve, 107.
furca Cleve var. macroceras Schroder,
107.
gracile Schutt, 79.
grunowii Schtitt, 104.
ikari Skv., 102.
javanicum Cleve, 100.
laciniosum Schtitt, 79.
lauderi Ralfs var., 101.
lorenzianum Grun., 100.
medium Schutt, 104.
messanense Castr., 107.
misumense Gran & Yendo, 101.
okamurai Ikari, 106.
ostenfeldii Cleve, 79.
pelagicum Cleve, 79.
peruvianum Brightw., 106.
procerum Schutt, 107.
protuberans Lauder, 103.
radians Schutt, 104.
ralfsii Cleve,, 106.
reichelti Hustedt, 102.
saltans Cleve, 106.
schiittii Cleve, 106.
septentrionale Oestrup, 79.
siamense Ostenf., 101.
skeleton Schroder, 105.
sociale Lauder, 78, 102.
spirillum Castr., 108.
tortissimum Gran., 105.
varians (Lauder) V. Heurck, 108.
sp., 80, 107.
Chaetosphaeria eximia Sacc, 501.
Champereia cumingiana, 488.
manillana, 488.
sp., 488.
Chrysophyllum oliviformis, 487.
Chytridiales, 517.
Cintractia axicola (Berk.) Cornu, 484.
Cissus sp., 486.
Citrus, 513.
aurantifolia, 510.
decumana, 510.
maxima, 510, 513, 522, 624, 527, 528.
nobilis, 503, 509-512, 516, 521. 524, 526.
sp., 485, 521, 527.
spp., 519, 523, 524.
Cladosporium herbarum L., 533.
lineolatum Sacc, 533.
Claoxylum sp., 488.
Clasterosporium maydicum Sacc, 538.
Claviciptae, 493.
Clerodendron fragrans, 482.
inermis, 480.
minahassae, 480.
sp., 485.
Climacodium biconcavum Cleve, 98.
Clitoria ternatea, 532.
Clypeosphaeria bakeriana Rehm, 506.
Clypeosphaeriacese, 505.
Coccomyces, 495.
dubius Rehm, 517.
quadratus (Schw. & Kze.) Karst. var.
philippinus Rehm, 517.
Cockroaches, 556, 575, 578, 583.
Cocos nucifera, 487, 495, 501, 502, 505, 510,
519, 521, 524, 529, 535.
Coexistent infection with yaws and syphilis,
177.
Coffea arabica, 479, 520.
excelsa, 491.
spp., 479, 520, 531.
Coix lachryma-jobi, 498, 509.
COLE, HOWARD IRVING, Causes of irri-
tation upon injection of iodized ethyl
esters of Hydnocar pus-group oils, 377.
Coleoptera, 759.
Coleosporiacese, 481.
Coleosporium exaci Syd., 481.
knoxiae Syd., 481
merrillii P. Henn., 481.
Colletotrichum arecae Syd., 527.
arecae Syd. forma setis perpaucis prae-
dita, 528.
euchroum Syd., 528.
falcatum Went., 528.
gloeosporioides Penz., 528.
lussoniense Sacc, 528.
nigrum Ell. & Hals., 528.
payayae (Henn.) Syd., 528.
Colocasia antiquorum, 519.
esculentum, 519.
Commelina, 483.
Composition of Philippine kapok-seed oil,
131 ; of Philippine peanut oil, 199.
770
Index
Coniosporium bambusae (Thuem. ft Bo lie)
Sacc, 633.
extremorum &yd., 533.
oryzinum Sacc, 533.
unilateral Sacc, & Peyr., 533.
vinosum (B. & C.) Sacc, 533.
Coniothyrium coffeae Henn., 520.
Conosia irrorata (Wied.)f 27.
Construction, bored-hole latrine equipment
and, 681.
Control of the common pink mealybug
Trionymus sacchari (Cockerell) of
sugar cane on Negros, 221.
COPELAND, EDWIN BINGHAM, New or
interesting oriental ferns, 209.
Cordus Schoenh., 415, 434, 440.
peguanus Senna, 388, 415, 440.
Corethron pelagicum Brun., 97.
Coriaria, 257, 259, 260.
intermedia Mats., 257-259, 263.
japonica A. Gray, 258, 259.
myrtifolia Linn., 258, 261.
ruscifolia Linn., 258.
Coriariacece, 257.
Cormopus Kolbe, 392, 396.
Corynelia clavata (L.) Sacc, 503.
Coryneliaceae, 503.
Corypha elata, 505, 525.
Coscinodiscus, 95.
concinnus W. Sm., 96.
excentricus Ehrenb., 96.
fragilissimus Grun., 96.
hyalinus Grun., 98.
mirificus, 96.
papuanus, 96.
radiatus Ehrenb., 96.
sol Wall., 97.
Croton tiglium, 525, 533.
CRUZ, AURELIO O., and AUGUSTUS P.
WEST, Composition of Philippine ka-
pok-seed oil, 131 ; Composition of Phil-
ippine peanut oil, 199.
Cryptocarya- sp., 491.
Cucumis sativus, 523.
Cucurbitariaceae, 503.
Cudrania javanica, 536.
Cuestis diffusa, 488.
Culex, 43.
fatigans, 656.
pipiens, 657, 659, 660, 663.
quinquefasciatus Say, 40, 657, 659, 660,
667.
sp., 40.
(Culex) pipiens Linn., 656.
(Culex) quinquefasciatus Say, 656.
Cyanopsis psoraleoides, 522.
Cyanotis axillaris, 483.
Cyathea bontocensis Copel., 209.
callosa, 211.
calocoma (Christ.) Copel., 210.
caudata, 211, 212, 515.
dementis, 210.
contaminans (Wall.) Copel., 210.
Cyathea — Continued.
dupaxensis Copel., 211.
edafioi Copel., 211.
halconensis, 212.
heteroloba, 210.
mearnsii, 212.
melanophlebia Copel., 212.
merillii Copel., 212.
pustulosa (Christ.) Copel., 212.
sqnanticosta Copel., 212.
Cyathus, 517.
Cyclophyllidea Braun, 546.
Cyclostemon sp., 526.
Cynodons dactylis, 499.
Cynomys ludovicianus, 564.
Cyperus compressus, 480.
Cyphagogus Parry, 392, 395-397, 483, 436.
buccatus Kin., 386, 392-394, 400, 43G.
eichhorni Kbm., 386, 393, 394, 400, 436.
gladiator Kin., 886, 393, 400, 436.
humilis Kin., 386, 393. 400.
longulus Senna, 386, 393, 400, 436.
modiglianii Senna, 386, 394, 396, 400,
436.
planifrons Kin., 386, 394, 400, 436.
silvanua Senna, 386, 394, 400, 436.
simulator Senna, 886, 394, 400, 436.
tabacicola Senna, 386, 395, 400, 436.
westwoodi Parry, 386, 395, 400, 486.
whitei Westw., 386, 395, 400, 436.
Cyrilla, 513.
Cysticercus fasciolaris Rudolphi. 538, 546.
sp., 546.
Cytospora aberran3 Sacc, 521.
palmicola B. & Cke., 521.
D
Daemonorops, 490, 502.
Dalbergia sp., 499.
Daldinia concentrica (Bolt.) Ces. & de Not.,
513.
concentrica var. microspora (Starb.)
Theiss., 513.
escholzii Ehr., 513.
Darwiniella, 494.
Datura alba, 521, 526.
Davainea formosana Akashi, 585.
madagascariensis Garrison, 549, 585.
madagascariensis (Davaine) Garrison,
548.
Davaineidse Fuhrmann, 548.
Davaineinse, 548.
Decay of wood in automobiles in the Tropics,
189.
Deeringia baccata, 481.
Dematiaceae, 530.
Dendrodochium lussonense Sacc, 535.
Dengue, notes on, 593.
virus, resistance of, 601.
Derris elliptica, 500.
philippinensis, 515.
sp., 500.
Index
771
Desmodium pulchellum, 486.
sinuosum, 523.
triflorum, 487.
Detonula schroederi Gran, 99.
Diaporthe citrincola Rehm, 510.
recondita Sacc, 510.
Diatoms, pelagic, of Korean strait of the Sea
of Japan, 95.
plankton, from Vladivostok Bay, 77.
Diatrypaceae, 511.
Diatrypella barleriae Syd., 511.
psidii Syd., 511.
Dichotomella areolata Sacc, 534.
Dictydium cancellatum Macb., 89.
Dictyothyriella muscosa Syd., 491.
Dictyotopterus Kin., 399, 433, 437.
philippinensis Kin., 386, 399, 400, 437.
pulcherrimus Kin., 386, 399, 400, 437.
Diderma globosum Pers., 88.
rugosum, 88.
rugosum Macb. var. aaiatica Skv., 85, 88.
spumarioides Fries, 85, 88.
Didymella caricae Tassi, 507.
eutypoides Rehm, 507,
lussoniensis Sacc, 507.
orchnodes Rehm, 507.
seriata Rehm, 507.
Didymium dubium, 86.
Didymosphaeria anisomera Sacc, 507.
caespitulosa Sacc, 507, 508.
inconspicaa Rehm, 507.
striatula Penz. & Sacc, 507, 508.
Didymotrichia, 502.
Die Brenthiden der Philippinen-Inseln, 383.
Diedickea singularis Syd., 527.
Digenea v. Beneden, 539.
Dillenia philippinensis, 488.
Dimerina graffii Syd., 485.
Dimerium tayabense Yates, 485.
Dimerocalyx longipes, 516.
Dinochloa, 490, 504, 508.
sp„ 500.
Diochus conicicallis Mots., 518.
Dioscorea, 483.
aculeata, 525.
alata, 483, 532.
esculenta, 483, 495, 499, 522-525, 529,
532, 533.
sp., 499.
spp., 524, 532, 533.
Diospyros discolor, 483.
sp., 536.
Diphasium, 209.
Diplodia artocarpi Sacc, 521.
artocarpina Sacc, 521.
caricae Sacc, 521.
circinans B. & Br., 521.
cococarpa Sacc, 521.
cococarpa var. malaccensis Tassi, 521.
crebra Sacc, 520.
daturae Sacc, 521.
durionis Sacc. & Syd., 521.
manihoti Sacc, 521.
Diplodia — Continued.
mori West., 522.
phaseolina Sacc, 522.
ricinicola Sacc, 522.
synedrellae Sacc, 522.
Diplodina degenerans Diedicke, 522.
Diptera, 129.
Philippine, 9.
Dipterocarpus grandiflorus Blco., 195.
spp., 195.
Discodothis lobata Syd., 515.
Distomata Zeder, 539.
Ditylium brightwellii (West) Grun., 77, 100.
sol V. Heurck, 100.
Diurus Pasc, 426, 435, 442.
furcillatus Gyll., 389, 426, 427, 442.
philippinicus SVmna, 389, 426, 427, 442.
samarensis Kin., 389, 426, 427, 442.
shelfordi Senna, 389, 426, 427, 442.
Dolichopeza Curt., 269-271.
i so lata Alex., 269.
(Mitopeza) longicornis (Brun.), 272.
(Mitopeza) nigromaculata (Edw.), 272.
(Mitopeza) nitidirostris (Edw.), 272.
(Mitopeza) rizalensis Alex., 271, 272.
(Nesopeza) angustaxillaris Alex., 272,
273.
(Nesopeza) melanosteraa Alex., 272.
Dolichos gibbosus, 518.
lablab, 507, 518, 526.
uniflorus, 507, 526.
Donax cannaeformis, 493, 505, 509.
Dothidea, 493, 494.
Dothideales, 494.
Dothidella, 500.
gigantochloae (Rehm) Theiss. & Syd.,
494.
Dothidiacese, 494.
Dothiorella crastophila Sacc, 522.
Dracontomelum cumingianum, 506.
Dryopteris clemensiae Copel., 218.
otaria, 213.
parasitica (L.) O. K., 213.
Dunbaria ferrignes, 518.
sp., 523.
Dang beetles, 578.
Durio zibethinus, 521, 525.
Dysoxylum, 524.
E
Earwig, 555.
Echinorhynchata Faust, 582.
Echinorhynchus cestodiformis Linstow, 582,
moniliformis Bremser, 582.
Echinostoma gotoi Ando, 544.
ilocanum, 539.
Echinostomatidse Looss, 539.
Echinostomatinae Looss, 539.
Echinostomatoidea Faust, 539.
Enterobius vermicularis, 570.
Ectocemus Pasc, 424, 441.
badeni Kbm., 389, 424, 441.
Elaeagnus philippinensis, 482.
772
Index
Ellisiodothis pandani Syd., 495.
rehmiana Theiss. & Syd., 495.
Elmerococcum orbicula Syd., 494.
Elsinoe canavaliae Rac, 517.
Elsinoeae, 517.
Endoxyla mangiferae Henn., 510.
Entyloma oryzae Syd., 484.
Ephydatia, 73.
fortis Weltner, 61, 71.
Epichloe warburgiana- P. Magn., 493.
Epigogus Kin., 396, 433, 436.
flexibilis Kin., 886, 396, 400, 436.
Epiphragma, 448.
bakeri Alex., 23.
ochrinota, 24.
(Folyphragma) angusticrenula Alex.,
468, 469.
(Polyphragma) apoensis Alex., 463, 466.
(Polyphragma) bakeri Alex., 23.
(Polyphragma) caninota Alex., 466, 468.
(Polyphragma) cinereinota Alex., 467.
(Polyphragma) crenulata Alex., 467-469.
(Polyphragma) fulvinota Alex., 461, 462,
464.
(Polyphragma) fnscofasciata Alex., 459,
460.
(Polyphragma) fuscosternata Alex., 464,
466.
(Polyphragma) griseicapilla Alex., 467,
468.
(Polyphragma) hastata Alex., 464, 466.
(Polyphragma) latitergata Alex., 460.
(Polyphragma) nigrotibiata Alex., 462.
(Polyphragma) ochrinota Alex., 23, 460,
462, 464.
(Polyphragma) parviloba Alex., 28.
Eriocera, 26.
Erioptera Meig., 286, 289.
argentifrons Edw., 287.
fusca de Meij., 287.
melanotaenia Alex., 287.
nigribaais Edw., 287.
parallela Brun., 287.
punctipennis Brun., 287.
subfusca Edw., 287.
(Empeda) gracilis (de Meij.), 289.
(Empeda) lunensis Alex., 288, 289.
(Empeda) rubripes Alex., 27.
(Teleneura) fusca de Meij., 287.
(Teleneura) melanotaenia Alex., 287.
EriopteTini, 28, 269, 284.
Erysiphaceae, 484.
Erythrina indica, 483, 534.
ESGUERRA, P. D., see Santos, West, and
Esguerra.
Esters, iodized ethyl, causes of irritation
upon injection of, of hydnocarpus-
group oils, 377.
Eterozemus Senna, 898, 433, 437.
hetus Senna, 386, 398, 400, 487.
pubens Senna, 386, 399, 400, 437.
Ethmodiscus convexus Castr., 96.
Eucampia, 95.
biconcava (Cleve) Ostenf., 97.
hemiauloides Ostenf., 97.
zodiacus Ehrenb., 97.
Euchlaena luxurians, 525.
Eugenia, 490, 508.
bataanensis, 506, 509.
calubcub, 490.
jambolana, 488.
Eunematoda Ward., 560.
Euparyphium Dietz, 539, 545.
guerreroi Tubangui, 538, 542, 584.
ilocanum (Garrison) Tubangui, 588, 589,
544, 584.
murinum Tubangui, 538, 543, 584.
Eupeithes Senna, 418, 435, 441.
dominator Kin., 388, 418, 422, 441.
Euphorbia neriifolia, 528.
Eupsalis Lac, 420, 421, 434, 441.
kleinei Heller, 389, 420, 422, 441.
Eutrachelini, 433.
Eutrixopsis javana Tns., 129.
Eutypa bambusina Penz. & Sacc, 510.
heteracantha Sacc, 510.
ludibunda Sacc, 510.
Eutypella citricola Speg., 510.
cocos Ferd. & Winge, 510.
leucaenae Rehm, 511.
lineolata Rehm, 511.
malloti Rehm, 511.
rehmiana (Henn. & Nym.) v. Hohn., 511.
Evonymus japonicus, 491.
Exacum chironioides, 481.
Exarmidium blumeanum (Rehm) Theiss. &
Syd., 498.
Exosporium durum Sacc, 535.
pulchellum Sacc, 535.
F
Fascioletta ilocana Garrison, 539.
Ferns, oriental, new or interesting, 209.
Fever, rat-bite, in the Philippines, 159.
Ficus, 507, 534.
banahaensis, 501.
carica Linn., 482, 483.
caudatifolia, 534.
garcia, 497.
hauili, 498.
heterophylla, 498, 501.
minahassae, 497, 517.
nervosa, 497.
odorata, 497.
odoratus, 497.
pseudopalma, 492.
ulmifolia, 489.
validicaudata, 497.
sp., 497, 498, 523.
Fimbristylis diphylla, 484.
Fleas, rat, 556.
FLEMING, WM. D., see Holt, Fleming,
and Kintneb.
Index
773
Fly Eutrixopsis javana Townsend (Diptera,
Tachinidse), a parasite of the beetle
Leucopholis irrorata in Occidental Ne-
gros, Philippine Islands, 129.
Freycinetia, 512.
Fuligo muscorum Alb. & Schwein, 85, 87.
septica Gmel., 85, 87.
septica Gmel. var. rufa R. E. Fries, 87.
Fumago vagans Pers., 488.
Fungi imperfecti, 520.
Fungi, second supplement to the list of
the lower, of the Philippine Islands,
479.
Fusarium cubense Efs., 535.
theobromae App. & Strunk., 536.
G
Ganguleterakis gangula Lane, 571.
GEE, N. GIST, Fresh-water sponges of the
Philippine Islands, 61.
Gibbera philippinensis Rehm, 503.
Gibberella saubinetii (Mont.) Sacc, 492.
Gigantochloa scribneriana, 494, 516.
Gigantorhynchus moniliformis (Bremser)
Railliet, 582.
Gilletiella, 491.
latemaculans Rehm, 490.
Gliricidia maculata, 510, 523.
sepium, 492, 526, 531.
Gloeosporium affine S'acc., 528.
alchorneae Syd., 528.
alstoniae Sacc, 528.
canavaliae Syd., 528.
catechu Syd., 528.
lebbek Syd., 528.
macrophomoides Sacc, 528, 529.
musarum Cke. & Mass., 529.
palmarum Oud., 529.
vanillae Cke., 529.
Glomerella cingulata (Stonem.) S. & v. S.,
509.
Glycine hispida, 481, 484.
javanica, 518.
max, 481, 484.
sojae, 481, 484.
Gmelia philippinensis, 485.
Gmelina, 488.
Gnomoniacese, 509.
Gongylonema Molin, 575.
neoplasticum (Fibiger & Ditlevsen) Ran-
som & Hall, 538, 575, 576, 584.
orientale Yokogawa, 575.
pulchrum, 576.
Gongyloneminse Hall, 575.
Goniothalamus, 507.
Gonomyia diffusa, 31.
flavomarginata (Brun.), 32, 33.
incompleta Brun., 29.
(Leiponeura) insulensis Alex., 29.
(Lipophleps) alboannulata Alex., 31, 32.
(Lipophleps) bicolorata Alex., 27.
Gonomyia — Continued.
(Lipophleps) diffusa (de Meij.), 32.
(Lipophleps) flavocostalis Alex., 33.
(Lipophleps) incompleta Brun., 29.
(Lipophleps) liberiensis Alex., 31.
(Lipophleps) luteimarginata Alex., 32.
(Lipophleps) maquilingia Alex., 28, 29.
(Lipophleps) noctabunda Alex., 31.
(Lipophleps) nubeculosa de Meij., 31.
(Lipophleps) pallidisignata Alex., 30.
(Lipophleps) robinsoni Edw., 34.
(Lipophleps) secreta Alex., 33, 34.
(Lipophleps) sobrina Alex., 31.
(Progonomyia) brunnescens Edw., 286.
(Progonomyia) tenebrosa Edw., 286.
(Progonomyia) terebrella Alex., 285, 286.
(Ptilostena) punctipennis Edw., 35.
Gossypium sp., 483.
spp., 481.
Grammatophora japonica Grun., 111.
marina (Lyngb.) Kutz., 112.
Grammitis bulbotricha, 219.
congener, 219.
fasciata, 220.
limapes Copel., 218, 219.
longa Fee, 220.
multifolia Copel., 219.
multifolia Copel., var. jg lasiosora Blm.,
220.
nana Fee, 220.
pubinervia, 219, 220.
pusilla, 219.
setosa, 220.
stenocrypta Copel., 220.
Graphiola arengae Rac, 536.
cylindrospora Syd., 536.
Grewia, 515.
stylocarpa, 506.
Guignardia arengae Rehm, 504.
bambusina Rehm, 504.
creberrima Syd., 504.
dinochloae Rehm, 504.
manihoti Sacc. var. diminuta S'acc., 504.
Guinardia baltica Schiitt, 114.
flaccida (Castr.) Per., 114.
Guioa perrottetii, 4S6.
Gymnema tingentis, 482.
Gymnospora spinosa, 486.
H
HADDEN, F. C, and A. W. LOPEZ,
Efforts toward biological control of the
common pink mealybug Trionymus
sacchari (Cockerell) of sugar cane
on Negros, 221.
Hadronema orbiculare Syd., 534.
Haemoproteus, 656.
Hamaspora acutissima Syd., 480.
Haplospora manilensis Sacc, 522.
Hare, European, 564.
Harknessia, 495.
774
Index
Heligmosominae Travassoe, 567.
Heligmosomum muris Yokogawa, 667.
Heliotropus indieus, 580.
Helius (Eurhamphidia abnormalis (Brun.),
283, 284.
(Eurhamphidia) diacanthus (Alex.), 283,
284.
(Eurhamphidia) fuscofemoratus Alex.,
282, 283.
(Eurhamphidia) indivisus Alex., 288.
(Eurhamphidia) nigrofemoratus (Alex.),
283.
(Helius) apoensis Alex., 455.
(Helius) arcuarius Alex., 455.
(Helius) procerus Alex., 454, 455.
(Helius) trianguliferus Alex., 456.
(Rhampholimnobia) reticularis (Alex.),
23.
Helminthosporium caryopsidum Saec, 584.
curvulum Sacc, 534.
ficinum Sacc, 534.
ficinum Yates, 534.
inconspicuum C. & Ell., 634.
inversum Sacc, 634.
oryzae Breda de Haan, 534.
papayae Syd., 534.
ravenelii Berk. & Curt., 584.
Helotiacese, 516.
Hemileia canthii Berk. & Br., 470.
vastatrix Berk. & Br., 479.
Hemisphaeriacese, 491.
Hemisphaeriales, 488.
Hemitrichia clavata Rost., 85, 91.
serpula Rost., 91.
vesparium Macbr., 91.
Henarrhenodes K. M. Heller, 423, 436, 441.
macgregori Heller, 389, 423, 441.
Henseniella baltica Schutt, 114.
Hepaticola Hall, 664.
hepatica (Bancroft) Hall, 688, 564, 566,
584.
Herpotrichia, 502.
Heterakidse Railliet & Henry, 671.
Heterakiiwe Railliet & Henry, 571.
Heterakis Dujardin, 571.
gallinae, 572.
spumosa Schn., 538, 571, 584.
Heteroblysmia Kin., 423, 585, 441.
accurata Kin., 389, 428, 424, 441.
electa Kin., 389, 423, 424, 441.
formidolosa Kin., 389, 423, 424, 441.
Heterodothis leptotheca Syd., 494.
Heteroplites Lac, 426, 442.
Pasc, 435.
erythroderes Boh., 389, 426, 427, 442.
Hevea, 619.
brasiliensis, 491, 494, 616, 522.
Hewittia sublobata, 486.
Hexatomini, 28, 457.
Heynea sumatrana, 489.
Hibiscus, double, somatic segregation in,
627.
esculentus, 492.
Hibiscus — Continued .
rosa-sinensis Linn., 628, 636.
sabdariffa, 524.
syriacus Linn., 633.
Higonius Lewis, 418, 434, 440.
cilo Lewis, 388, 418, 417, 440.
crux, 413.
Histamine test as an aid in the diagnosis
of early leprosy, 123.
Holcus sorghum, 480, 484, 488, 500, 507,
534.
HOLT, R. L., and J. H. KINTNER, Notes
on Dengue, 593.
HOLT, R. L„ WM. D. FLEMING, and J.
H. KINTNER, Resistance of dengue
virus, 601.
HomophyhiB Kin., 406, 434, 438.
mindanensis Kin., 387, 406, 416, 438.
Hoplopisthius Senna, 414, 400, 434.
trichimerus Senna, 388, 414, 417, 440.
Hormocerus Schoenh., 428, 435, 442.
reticulatus F., 389, 428, 432, 442.
scrobicollis, 428.
Hormorhynchus moniliformis (Bremser)
Ward, 582.
Hoya, 528.
luzonica, 508.
Humaria caballina Rehm, 516.
HUMPHREY, C. J., Decay of wood in au-
tomobiles in the Tropics, 189.
Hymenolepidida* Railliet & Henry, 558.
Hymenolepidinae Ransom, 553.
Hymenolepis Weinland, 658.
diminuta (Rudolphi) Blanch., 538, 568,
584.
flavopunctata Weinland, 558.
fraterna Stiles, 556.
longior Bay lis, 666.
nana (Sieb.) Blanch., 554, 556, 584.
vide Joyeux, 556.
vide Woodland, 556.
Hymenopsis cudraniae Mass., 686.
Hymenula copelandi Sacc, 536.
Hyphales, 530.
Hypocrea, 498.
Hypocreaceae, 491.
Hypocreales, 491.
Hypocrella cretacea, 403.
discoidea (Berk. & Br.) Sacc, 498, 627.
globosa, 493.
grewiae, 493.
mollii Koord., 493, 626.
pernettyae, 493.
reciborskii A. Zimm., 493.
reineckiana P. Henn., 493, 527.
salaccensis (Rac) Petch., 498.
schizostochyii P. Henn., 498.
warneckiana, 493.
zimmermanniana, 493.
zingiberis, 493.
Hypodermataceae, 514.
Index
775
Hypomiolispa Kin., 411, 412, 484, 489.
exarata Desbr., 888, 411, 416 489.
helleri Kin., 888, 411, 416, 440.
nupta Senna, 388, 411. 412, 417, 440.
ocularis Kin., 888, 412, 418, 440.
sponsa Kin., 388, 412, 417, 440.
tomentosa Kin., 388, 412, 417, 440.
trachelizoides Senna, 388, 412, 417, 440.
Hypoxylon annulatum (Schw.) Mont., 512.
atropurpureum Fr., 512.
culmorum Cke., 512, 518.
effusum Nitsch., 512.
freycinetiae Rehm, 512.
granulosum Bull., 512.
haematostroma Mont., 512.
marginatum (Schw.) Berk., 512.
marginatum (Schw.) Berk. var. mammi-
forme Rehm, 512.
rubigineo-areolatum Rehm var. microi-
porum TheiBS., 512.
subeffusum Speg., 512.
Hyptis suaveolens, 486.
Hysteriacese, 515.
Hysteriales, 514.
Hysterium, 514.
anceps Sacc, 515.
HyBterostomella, 495, 515.
latracerae (Rud.) v. Hoehn., 495.
spurcaria (Berk. & Br.) v. Hoehn., 495.
tetracerae (Rud.) v. Hoehn., 495.
Idiophlebia Grunberg, 294.
Illosporium tabacinum Sacc, 586.
Imperata cylindrica, 492, 504, 585.
Inocyclus psychotriae Syd., 495.
Interpretation of the laws of Brown and
Pearce that govern the course of tre-
ponematoses, 169.
Ipomoea batatas, 522.
pes-caprae, 482.
Isachne miliacea, 484.
Ischaemum ariitatum, 484.
Isothea, 499.
Itea macrophylla, 486.
Ithyetenini, 390, 425, 427, 482, 433, 485,
442.
Ixora sp., 487.
JonthoceruB Lac, 899, 400, 434, 437.
asiaticus Kin., 387, 399, 401, 405, 487.
bicolor Heller, 387, 400, 405, 487.
laticostatis Kin., 387, 401, 405, 437.
modiglianii Senna, 387, 401, 405, 437:
Justicia gendarussa, 480.
Kahn test in clinical syphilis, 225.
Kapok-seed oil, composition of, 181.
KING, W. V., The Philippine varieties of
Anopheles gigas and Anopheles linde-
sayi, 751.
KINTNER, J. H., see Holt and Kintnbr ;
see also Holt, Fleming, and Kint-
nbr.
KLEINE, R., Die Brenthiden der Philip-
pinen-Inseln, 888.
Knoxia corymbosa, 481.
Korean strait of the Sea of Japan, pelagic
diatoms of, 95.
Kotlania madagascariensis (Davaine), 586.
Kretzmaria ghomphoidea Penz. & Sacc, 518.
Kuehneola desmium (Berk. & Br.) Arth.,
481.
fici (Cast.) Butl., 482.
fici (Cast.) Butl. f. moricola F. Henn.,
482.
Labia sp., 518.
Lachnea livida (Schum.) Gill., 516.
lurida P. Henn. & E. Nym., 516.
Lactuca sativa, 581.
Lagenaria leucantha, 509.
Lagerheima dermatoidea Rehm, 515.
Lagerstroemia indica, 515, 517.
speciosa, 515, 517.
Laosa, 448.
Lasiodiplodia theobromae (Pat.) Griff. &
Maubl., 522.
Lasiosphaeria mollis Rehm, 501.
Lasiothyrium cycloschizon Syd., 527.
Latrine equipment and construction, 681.
Lauderia, 95.
annulata Cleve, 99.
borealis Gran, 99.
compressa Per., 99.
delicatula Per., 99.
Lauderiopsis costata Ostenf., 99.
Laws of Brown and Pearce, an interpreta-
tion of the, that govern the course
of treponematoses, 169.
Lawsonia inermis, 489.
Leaf and seed structure of a Philippine
Coriaria, 267.
Lembosia, 496.
congregata Syd., 490.
crustacea (Cke.) Theiss., 490.
eugeniae Rehm, 490.
javanica (Pat.) Rac, 490.
pandani (Rostr.) Theiss., 490.
pothoidei Rehm, 490.
Lennites striata, 193.
LEON, WALFRIDO DE» see Oliver, Lion,
and Roda.
Lepidiota pruinosa, 761.
Lepidium sativum, 518.
Lepidoderma mandshurica Skv., 85, 88.
Lepisanthes schizolepis, 491.
Lepistemon fiavescens, 482.
Leprosy, histamine test as an aid in the
diagnosis of early, 128.
776
Index
Leptamorphocephalus Kin., 415, 434, 440.
fcederatus Kin., 888, 415, 440.
Leptochilus lanceolatus, 220.
Leptocylindrus curvatus Skv., 100.
danicus Cleve, 77.
Leptosphaeria orthogramma (B. & C.) Sacc,
508.
Leptostromatacese, 527.
Leptothyrium cireumscissum Syd., 527.
Lepus europus, 564.
Leucaena glauca, 511.
Leucopholis irrorata Chevr., 129, 759.
Libnotes, 448, 449, 451.
Licea biforis Morgan, 86, 89.
brassica Skv., 85, 89.
flexuosa Pers., 89.
mandshurica Skv., 85, 89.
Limacinia biseptata Sacc, 488.
Limacinula malloti Rehm, 488.
Limnophila granulata, 25.
(Ephelia) granulata Edw., 25.
(Ephelia) igorota Alex., 24, 25.
Limonia, 449, 451.
argentifera de Meij., 19, 20.
cynotis, 14, 15.
manca Alex., 19, 20.
morionella Schiner, 16.
multinodulosa, 12.
nigronitida Alex., 20.
nigronotata Brun., 20.
pleuropalloris Alex., 20.
rostrifera, 16.
sorbillans (Wied.), 20.
unicinctipes, 21.
(Alexandriaria) sollicita Alex., 21, 22.
(Dicranomyia) fullowayi (Alex.), 282.
(Dicranomyia) mesosternata (Alex.), 281.
(Dicranomyia) mesosternatoides (Alex.),
281.
(Dicranomyia) neopunctulata Alex., 282.
(Dicranomyia) orthia Alex., 281.
(Dicranomyia) punctalata (de Meij.),
282.
(Dicranomyia) sordida (Brun.), 22.
(Dicranomyia) subpunctulata Alex., 282.
(Geranomyia) argentifera (de Meij.), 27.
(Geranomyia) longifimbriata Alex., 18,
19.
(Geranomyia) paramanca Alex., 19, 20.
(Geranomyia) phoenosoma Alex., 17, 18.
(Goniodineura) nigriceps, (van der
Wulp), 27.
(Laosa) gloriosa (Edw.), 448.
(Laos a) manobo Alex., 447-449.
(Libnotes) amatrix Alex., 280.
(Libnotes) klossi Alex., 280.
(Libnotes) melancholic* Alex., 278, 279.
(Libnotes) neofamiliaris Alex., 277, 279.
(Libnotes) perrara Alex., 279, 280.
(Libnotes) subfamiliaris Alex., 277, 279.
(Libnotes) terrae-reginas Alex., 280.
(Libnotes) unistriolata Alex., 276, 279.
(Limnobia), 16, 282.
Limonia — Continued.
(Limonia) atroaurata Alex., 450.
(Limonia) bagobo Alex., 452.
(Limonia) bilan Alex., 449, 450.
(Limonia) bilobulifera Alex., 273, 274.
(Limonia) candidella Alex., 11, 12.
(Limonia) canis Alex., 14, 15, 452.
(Limonia) cynotis Alex., 452.
(Limonia) flavohumeralis Alex., 13, 14.
(Limonia) latiflava Alex., 12, 13.
(Limonia) luteivittata Alex., 276.
(Limonia) melanopleura Alex., 274, 275.
(Limonia) pacata* Alex., 453.
(Limonia) prolixicornis Alex., 453, 454.
(Limonia) retrusa Alex., 14.
(Limonia) subpacata Alex., 453.
(Limonia) subprolixa Alex., 453, 454.
(Limonia) tremula Alex., 275, 276.
(Pseudoglochina) angustapicalis Alex.,
20, 21.
(Pseudoglocbina) unicinctipes Alex., 21.
(Rhipidia) luteipleuralis Alex., 16.
(Rhipidia) morionella (Edw.), 15, 16.
(Thrypticomyia) apicalis (Wied.), 27.
Limoniinse, 11, 273, 447.
Limoniini, 11, 273, 447.
Lindsaya apoensis, 216.
longa Copel., 216.
macraeana, 216.
merrillii, 216.
protracta, 216.
Linospora elasticae Koord., 507.
pandani Rehm, 507.
seriata (Syd.) Rehm, 507.
Lipophleps, 31.
Lisea revocans Sacc, 492.
Litsea glutinosa, 486, 531.
mollis, 486.
Livistona, 514, 536.
Lomariopsis smithii Fee, 216.
LOPEZ, A. W., The fly Eutrixopsis javana
Townsend (Diptera, Tachinidae), a
parasite of the beetle Leucopholis irro-
rata in Occidental Negros, Philippine
Islands, 129 ; The use of the antennae
as a means of determining the sexes
in Leucopholis irrorata adults (Coleop-
tera, Scarabasidae), 759; see also H ad-
den and Lopez.
Lophodermium aleuritis Rehm, 514.
arundinaceum (Schrad.) Chev., 514.
arundinaceum (Schrad.) Chev. f. vul-
gare Fckl., 514.
passiflorae Rehm, 514.
planchoniae Rehm, 515.
rotundatum Syd., 515.
Loranthomyces sordidula (Lev.) v. Hoehn.,
490.
Loranthus haenkeani, 490.
sp., 490.
Lucidium pythiodes, 518.
Lycogala epidendrum Fries, 85, 90.
Lycoperaicum esculentum, 518, 519, 536.
Index
111
Lycopodium delbrueckii, 209.
edanoi Copel., 209.
phlegmarioides, 209.
M
Macaranga, 636.
tanarius, 486.
utilis, 529.
sp., 488, 529.
Macrophoma arengae Sacc, 522.
cyanopsidis Syd., 522.
musae (Cke.) Berl. & Vogl., 522.
obsoleta Sacc, 523.
trichosanthis Syd., 523.
Malaria, avian, 305, 347, 651.
transmission in the Philippines, III, 47 ;
IV, 247 ; V, 363 ; VI, 371.
Mallotus philippinensis, 488, 511.
Malpighia, 506.
MANALANG, C, Origin of the irritating
substance in mosquito bite, 39 ; Mal-
aria transmission in the Philippines,
III, 47; IV, 247; V, 363; VI, 371.
Manchuria, China, Mycetozoa from north, 85.
Mangifera indica, 486, 509, 510, 527, 529, 532.
Manihot dichotoma, 525.
utilissima Pohl, 504, 521, 524, 528, 531,
532, 627.
Marchalia, 495.
constellata (Berk. & Br.) Sacc, 495.
spurcaria, 495.
Marsonia pavonina Syd., 529.
Massalongiella imperatae Rehm, 504.
Massaria bataanensis Rehm, 509.
Massariacese, 509.
Massarina raimundoi Rehm, 509.
Massarinula bambusicola Rehm, 509.
donacina Rehm, 509.
obliqua Sacc, 509.
Mealybug, pink, efforts toward biological
control of the common, of sugar cane
on Negros, 221.
Megalonectria pseudotrichia (S'chw.) Speg.,
491.
Megistomastix Alex., 270.
nigromaculata Edw., 270.
Melampsoracese, 481.
Melanconiaceae, 527.
Melanconiales, 527.
Melanconidaceae, 511.
Melanconium sacchari Cke., 529.
Melanomma mindorense Rehm, 501.
Melanopsamma lichenoides Rehm, 501.
Melastoma, 527.
fusca, 498.
Melia azedarach, 533.
Meliola affinis Syd., 485.
aliena Syd., 485.
alstoniae Koord., 485.
apayaoensis Yates, 486.
arachnoidea Speg., 485.
arundinis Pat., 485.
Meliola — Continued.
bakeri Syd., 485.
callicarpae Syd., 485, 586.
callista Rehm, 485.
citricola Syd., 485.
clerodendricola P. Henn., 485.
cookeana Speg. var. saccardoi Syd., 486.
cylindrophora Rehm, 486.
desmodii Karst. & Roum., 486.
dichotoma Berk. & Cke., 486.
elmeri Syd., 486.
gymnosporiae Syd., 486.
hewittiae Rehm., 486.
hyptidis Syd., 486.
intricata Syd., 486.
macarangae Syd., 486.
maesae, 492.
mangiferae Earle, 486.
merremiae Rehm, 486.
merrillii Syd., 486.
micromera Syd., 485.
mitragynes Syd., 486.
panici Earle, 486.
parenchymatica Gaill., 487.
perpusilla Syd., 487.
piperina Syd., 487.
polytricha Kalch. & Cke., 487.
quadrispina Rac, 487.
sandorici Rehm, 487.
sidae Rehm, 487.
substenospora v. Hoehn. f. rottboelliae
Rehm, 487.
tamarindi Syd., 487.
telosmae Rehm, 487.
uncariae Rehm, 487.
sp., 489.
Melogrammatacese, 511.
Memecylon lanceolatum, 489.
subfurfuraceum, 489.
sp., 485.
MENDIOLA, N. B., Somatic segregation in
double Hibiscus and its inheritance,
627.
Merremia hederacea, 486.
umbellata, 487.
sp., 489.
Merrilliopeltis calami P. Henn., 490.
daemonoropsis Syd., 490.
hoehnelii Rehm, 490.
Mesoderes Senna, 398, 433, 437.
fessus Kin., 386, 398, 400, 437.
Mesomyites Ckll., 294.
Metasphaeria corruscans Rehm, 508.
incomplete- Rehm, 508.
maculans, 501.
Metatrachelizus Kin., 406, 484, 488.
constans Kin., 387, 406, 416, 438.
Microdiplodia passeriniana (Thum.) A 11 each.,
523.
Microdothella culmicola Syd., 495.
Micropeltella consimilis Rehm, 491.
Micropeltis aeruginascens Rehm, 491.
mucosa Syd., 491.
26421
16
778
Index
Microthyriaceae, 488.
Microthyriella latemaculans (Rehm) Theiss.
& Syd., 491.
philippinensis Syd., 491.
Microthyrium, 496.
Microtrachelizus Senna, 413, 414, 434, 440.
flexus Kin., 417.
fluxus Kin., 388, 413, 440.
pubescens Senna, 388, 413, 417, 440.
siamensis Kin., 38, 413, 417, 440.
tabaci Senna, 388, 413, 417, 440.
Microxyphium dubium Sacc, 488.
Milletia, 499.
cavitensis, 499.
Miolispa Pasc, 407, 410, 412, 431, 434, 438.
bicolor Kin., 387, 407, 416, 439.
clavicornis Kin., 387, 407, 416, 439.
cruciata Senna, 387, 407, 416, 439.
discors Senna, 387, 407, 416, 439.
elongata Kin., 387, 408, 410, 416, 439.
ephippium Kin., 387, 408, 416, 489.
flavolineata Kin., 387, 408, 416, 489.
flexilis Kin., 387, 408, 416, 439.
formosa Kin., 387, 408, 416, 439.
fornicata Kin., 387, 408, 416, 439.
fraudatrix Kin., 387, 409, 416, 488.
intermedia Senna, 387, 409, 416, 489.
lineata Senna, 387, 409, 416, 439.
pascoei Kin., 387, 409, 416, 439.
paueicostata Kin., 387, 409, 416, 488.
persersimilis Kin., 388.
persimilis Kin., 409, 416, 438.
pulchella Kin., 388, 410, 416, 488.
robusta Kin., 388, 410, 416, 489.
siporana Senna, 388, 410, 416, 439.
unicolor Kin., 388, 410, 416, 438.
Miscanthus japonicus, 514.
Mischocarpus fuscescens, 509, 535.
Mitopeza Edw., 270.
Jongicornis Brim., 270, 272.
mjobergi Edw., 270, 272.
nigromaculata (Edw.), 272.
nitidirostris (Edw.), 270, 272.
rizalensis Alex., 270, 272.
Mitragyne rotundifolia, 486.
Mollisia ravida Syd., 515.
Mollisiaceae, 515.
Molophilus banahaoensis Alex., 289, 290.
kempi Alex., 289, 290.
mendieus Alex., 292, 293.
procericornis Alex., 290, 292.
tawagensis Alex., 293, 294.
Momordica sp., 485.
Moniliformidae Van Cleave, 582.
Moniliformis Travassos, 582.
moniliformis (Bremser) Travassos, 582,
584.
MONSERRAT, CARLOS, The Kahn test in
clinical syphilis, 225 ; Comparative se-
rologic study of Vernes, Wassermann,
and Kahn reactions in experimental
treponematoses, 241.
Morenoella breviuscula, 490.
memecyli Syd., 489.
Morus alba, 482, 484, 511, 520, 522, 526.
albus, 482.
Mosquito bite, origin of the irritating sub-
stance in, 39.
Mosquitoes, Anopheles, daytime resting places
of, in the Philippines, 639.
Moth, meal, 555.
Mucedinaceae, 530.
Mucoraceas, 520.
Mucuna deeringiana, 481, 532.
nivea, 481.
Munkiella, 498.
Munkiodothis melastomata (v. Hoehn.)
Theiss. & Syd., 498.
Mus decumanus Pallas, 537.
norvegicus Erxleben, 587, 569, 584.
Musa cavendishii, 504.
paradisiaca sapientum, 504, 516, 523,
535.
sapientum, 504, 516, 521, 522, 529, 585.
textilis, 504, 520, 528, 530, 535.
sp., 521.
Mycelia sterilia, 536.
Mycetozoa from north Manchuria, China,
85.
Mycobacterium leprae, 128, 611-615, 616-623.
Mycochytridiaceae, 518.
Mycogne cervina Ditm. var. theobromae
Sacc., 530.
Mycorales, 520.
Mycosphaerella aloeasiae Syd., 504.
aristolochiae Syd., 504.
brideliae Syd., 504.
caricae Syd., 504.
musae S*peg., 504.
oculata Syd., 504.
pericampyli Syd., 505.
reyesii Syd., 505.
Mycosphaerellacese, 503.
Myiocoprella bakeri Sacc, 491.
Myiocopron bakerianum Rehm, 491.
Myocopron, 497.
Myriangiales, 517.
Myriangium duriaei Mont., 517.
N
Navicula kariana Grun. var. minor Grun.
forma curta Cleve, 113.
kariana Grun. var. minor Grun. forma
japonica Skv., 113.
pellucida Karst., 112.
(Cistula) lorenziana Grun., 112.
(Schizonema) mollis W. Sm„ 113.
(Schizonema) ramosissima Agardh for-
ma amplia Grun., 113.
Nectria bainii Mass., 492.
bainii Mass. var. hypoleuca Sacc, 492.
discophora Mont., 492.
striatospora, 492.
subfurfuracea P. Henn. & Nym., 482.
tjibodensis Penz. & Sacc. var. gliricidiae
Rehm, 492.
Nectriaceae, 491.
Index
779
Nectrioidaeeae, 526.
Nemathelminth.es Vogt, 560.
Nematoda Rudolphi, emend. Diesing, 560.
Nematode, 585.
Nematodes, 538, 684.
Nemocephalini, 433.
Neolitsea, 517.
Neopeckia rhodosticta (B. & Br.) Sacc, 502.
rhodosticta (Berk. & Br.) Sacc. var.
magnifica Rehm, 502.
Nesopeza Alex., 269, 270.
gracilis, 269.
Nicotiana tabacum, 518, 532, 536.
Nipa fruticans, 490, 496, 509.
Nippostrongylus Lane, 567.
muris (Yokogawa) Lane, 538, 560, 567,
584.
Niptera grewiae Rehm, 515.
Nitschkea bambusarum Rehm, 508.
Nitzehiella longissima (Breb.) Ralfs forma
parva V. Heurck, 81, 115.
Nitzschia birostrata Sm., 115.
Nummularia citrincola Rehm, 513.
fragillima Rehm, 513.
glycyrrhiza (B. & C.) Sacc, 518.
lianae Rehm, 513.
memorabilis Rehm, 513.
papyracea Rehm, 513.
reyesiana- Rehm, 513.
seutata B. & C, 513.
urceolata Rehm, 513.
O
Oidium erysiphoides Fr., 530.
Oil, kapok-seed, composition of, 131.
Philippine peanut, composition of, 199.
Philippine pine-needle, from Pinus in-
sularis (Endlicher), 1.
Oils, hydnocarpus-group, causes of irrita-
tion upon injection of iodized ethyl
esters of, 377.
Oleandra benguetensis Copel., 217.
chinensis Hance, 218.
colubrina, 217.
cumingii Presl., 217, 218.
macrocarpa- Presl., 217, 218.
maquilingensis, 217.
mollis, 217.
neriiformis, 217.
scandens Copel., 217, 218.
whitneii, 217.
OLIVER, WADE W., WALFRIDO DE
LEON, and ALFREDO PIO DE
RODA, The attempted cultivation of
Mycobacterium lepra?, 611.
Oomycetes, 517.
Oospora candidula Sacc, 530.
hyalinula Sacc. var. sordidula Sacc,
530.
oryzetorum Sacc, 680.
Operculina turpethum, 483.
Ophiobolus, 507.
heterostrophus Drechsler, 509.
nipae Henn., 509.
oryzae I. Miyake, 509.
oryzinus Sacc, 509.
Ophiochaete bakeriana Sacc, 508.
Ophiodothis thanatospora (Lev.) Rac, 493.
Ophionectria erinacea Rehm, 492.
theobromae (Pat.) Duss., 492.
Opisthenoplus Kin., 427, 435, 442.
ealabresii Kin., 389, 427, 432, 442.
cavis F. Walk., 389, 427, 428, 432, 442.
fascinatus Kin., 389, 428, 432, 442.
fecundus Kin., 389, 428, 432, 442.
madens Lac, 389, 428, 432, 442.
Opisthenoxys Kin., 397, 433, 437.
boettcheri Kin., 386, 397, 400, 437.
Orania palindan, 535.
Orchidaceae, 481, 491, 529.
Origin of the irritating substance in mog-
quito bite, 39.
Orimarga rubricolor, 23.
(Orimarga) rubricolor Alex., 22.
Oropeza Needham, 270.
Orthopareia Kin., 396, 433, 436.
idonea Kin., 386, 396, 400, 436.
Oryza sativa, 484, 494, 501, 509, 518, 525,
530, 533, 534, 536.
Oxyuridae Cobbold, 569.
Oxyuroidea Railliet, 569.
Ozonium glumicola Sacc, 536.
Pachypatella alsophilae (Rac) Theiss. &
Syd,. 515.
Pachyrrhizus angulatus, 482.
erosus, 482.
Paederia foetida, 483.
tomentosa, 483.
Pahudia rhomboidea, 499, 532.
Palawania cocoes Syd., 495.
grandis (Niessl.) Syd., 496.
Palmularia, 496.
Pandanus, 490, 502, 506.
copelandi, 490.
luzonensis, 495.
radicans, 506.
sabutan, 502, 505, 507.
tectorius, 494.
utilissima, 507.
Panicum, 481, 492.
auritum, 534.
indicum, 484.
repens, 481.
sp., 492, 494.
Paramorphocephalus Kin., 418, 434, 440.
setosus Kin., 388, 418, 440.
Paranectria luxurians Rehm, 492.
Parasite of the beetle Leucopholis irrorata
in Occidental Negros, Philippine Is-
lands, 129..
780
Index
Parasites, worm, of the brown rat (Mus
norvegicus) in the Philippine Islands,
537.
Parkia javanica, 499.
timoriana, 499.
Parodiella grammodes (Kze.) Cke., 487.
Passifiora quadrangularis, 491, 526.
Patellariacese, 515.
Pavetta- indica, 482.
Payena leeri, 509.
Pazschkiella philippinensis Yates, 523.
Peanut oil, composition of, 199.
Pelagic diatoms of Korean strait of the
Sea of Japan, 95.
Pennisetum, 499.
Peragallia meridiana Stohutt, 106.
Perieampylus incanus, 505.
Perichaena depressa Libert, 85, 92.
Periconia philippinensis Saco., 584.
Periplaneta americana, 578, 583.
Perisporiaceas, 485.
Perisporiales, 484.
Peroneutypella arecae Syd., 510.
graphidioides Syd., 510.
Peronospora, 519.
Peronosporaceas, 519.
Peronosporales, 519.
Persea americana, 509, 536.
gratissima, 536.
Pestalozzia funerea Desm., 529.
palmarum Cke. & Grev., 529.
pauciseta Sacc, 529.
Pezizaceae, 516.
Pezizales, 515.
Pezizella ombrophilacea Kehm, 516.
Phacidiaceae, 517.
Phacidiales, 517.
Phacidium, 495.
Phakospora pachyrhizi Syd., 482.
phyllanthi Diet., 482.
Phaodothis gigantochloae, 508.
Phaseolus aureus, 531.
calcaratus, 499.
lunatus, 522, 532, 583.
mungo, 481, 484.
vulgaris, 520, 522.
sp., 499, 531.
spp., 481, 508, 517, 528, 523.
Phellestroma hypoxyloides Syd., 524.
Phenacuspis mischocarpi, 535.
Philippine Anopheles gigas and Anopheles
lindesayi, varieties of, 751.
Coriaria, 257.
fresh-water sponges, 61.
kapok-seed oil, 131.
peanut oil, composition of, 199.
pine-needle oil from Pinus insularis
(Endlicher), 1.
rat-bite fever, 159.
Tipulidse, 9, 269.
Phoma bakeriana Sacc, 528.
citricarpa McAlpine, 523.
herbarum Westd., 524.
Phoma- — Continued.
musae Carpenter, 522.
oleracea Sacc, 524.
sabdariffae Sacc, 524.
sesamina Sacc, 524.
solanophila Oud., 524.
Phomatospora migrans Rehm, 510.
Phomopsis arecae Syd., 524.
calanthes Sacc, 523.
capsici (Magnaghi) Sacc, 523.
cinerescens (Sacc) Bubak, 523.
dioscoreae Sacc, 523.
gliricidiae Syd., 528.
palmicola (Wint.) Sacc. f. arecae Sacc,
524.
Phragmitis karka, 486.
vulgaris, 485.
Phycomycetes, 517.
Phyllachora, 495, 497, 498.
afzeliae Syd., 498.
canari P. Henn., 498.
circinata var. sanguinea, 498.
coicis P. Henn., 498.
cynodontis (Sacc.) Niessl., 499.
dalbergiae Niessl., 499.
dioscorea Schw., 499.
fici-fulvae, 497.
flci-minahassae, 497.
ficuum, 501.
luzoniensis P. Henn., 499.
minutissima (Welw. & Curr.) Sm., 499.
orbicula Rehm, 499.
pahudiae Syd., 499.
parkiae P. Henn., 499.
phaseolina Syd., 499.
pongamiae (Berk. & Br.) Petch., 499.
pterocarpi non Rehm, 498.
pterocarpi Rehm non Syd., 500.
rehmiana Theiss. & Syd., 499.
rottboelliae Syd. & Butl., 499.
roureae Syd., 499.
sacchari P. Henn., 500.
sacchari-spontanei Syd., 500.
sorghi v. Hoehn., 500.
spinifera, 497.
tjankorreh Rac, 500.
yapensis (P. Henn.) Syd., 500.
Phyllachoracese, 496.
Phyllactinia suffulta (Rebent.) Sacc, 484.
Phyllanthus niruri, 482.
reticulatus, 488.
sp., 482.
Phyllodromia germanica, 556, 575, 578.
Phylloporina phyllogena Muel.-Arg., 494.
Phyllosticta circumsepta Sacc, 524.
cocophylla Pass., 524.
densissima Sacc, 524.
dysoxyli Sacc, 524.
euchlaenae Sacc, 525.
glumarum Sacc, 525.
graffiana Sacc, 525.
insularum Sacc, 525.
manhoticola Syd., 525.
miurai I. Miyake, 525.
Index
781
Physalospora, 497.
affinis Sacc, 608.
bambusae (Rabh.) Sacc, 508.
bambusicola Rehm, 508.
dinochloae Rehm, 508.
guignardioides Sacc, 508.
hoyae v. Hoehn., 508.
peribambusina Rehm, 509.
Physarum, 86.
asiaticum Skv., 85, 87.
compressum Skv., 85, 86.
griseum Skv., 85, 86.
mandshuricum Skv., 85, 87.
Phytophthora colocasiae Rac, 519.
faberi Maubl., 519.
infestans (Mont.) de Bary, 619.
melongenae K. Sawada, 519.
phaseoli Thaxter, 519.
theobromae, 519.
Picea vulgaris, 4.
Pilaria, 27.
alboposticata Alex., 28.
carbonipes Alex., 27, 28.
carbonipes holomelania Alex., 28.
melanota Alex., 28.
phoenosoma Alex., 25, 26.
Pilobolaceae, 520.
Pilobolus lentiger Cda., 520.
Pilobus kleinii var. sphaerospora, 520.
Pilocratera tricholoma (Mont.) P. Henn.,
516.
Pinanga, 488.
Pine-needle oil, Philippine, from Pinus in-
sularis (Endlicher), 1.
Pinus contorta, 4.
excelsa, 4.
halepensis, 4.
insularis EndL, 1, 4.
lambertiana, 4.
longifolia, 4.
ponderosa, 4.
pumilis, 4.
sabiniana, 4.
sylvestris, 4.
Pionnotes capillacea Sacc, 536.
Piper retrofractum, 485.
sp., 487.
Pipturus arborescens, 489, 502.
Pittosporum pentandrum, 486.
sp., 486.
Placosphaeria duriones Syd., 525.
tiglii Henn., 525.
Placostroma pterocarpi (Mass.) Theiss. &
Syd., 500.
Planchonia spectabilis, 515.
Plankton diatoms from Vladivostok Bay, 77.
Planktoniella sol (Wall.) Schutt, 97.
PLANTILLA, FIDEL C, see Rodriguez and
Plantilla.
Plasmochin, prophylactic, in inoculated avian
malaria, 305.
prophylactic, versus prophylactic quinine
in inoculated avian malaria, 347.
Plasmodium cathemerium Hartm., 306, 808,
334, 347, 360, 656, 657.
Platyhelminthes Claus, 539.
Plectronia, 479.
didyma, 485.
horrida, 479.
Pleosporaceae, 507.
Pleurosigma arcuatum Donk., 81.
balticum var. wansbeckii Donk., 113.
fasciola Ehreb. var. arcuatum Donk., 81.
longum Cleve var. infiata Per. forma
japonica Skv., 113.
wansbeckii Donk., 113.
Plicaria bananincola; Rehm, 516.
tropica Rehm, 516.
Podocarpus, 503.
costatus, 503.
Polygonum chinensis, 480.
tomentosum, 480.
Polyosma philippinensis, 527.
sorsogonensis, 527.
Polyphragma, 467.
Polypodium alpestre Blm., 219.
bulbotrichum Copel., 219.
Polyporus, 516.
sanguineus, 193.
Polyscias nodosa, 512.
Polystomellaceas, 494.
Pongamia glabra, 499.
mitis, 499.
pinnata, 499.
Popillia japonica, 129.
Populus simonii, 92.
Pothoideum lobbiamim, 490.
Prairie dog, 564.
Premna, 516.
cumingiana, 484, 507, 508.
odorata, 485, 493, 504, 507, 515.
vestita, 484.
sp., 489, 504.
Prophthalmus Lac, 419, 435, 441.
longirostris Gyll., 388, 419, 422, 441.
tricolor Power, 388, 419, 422, 441.
Propolidiopsis arenga Rehm, 517.
Prosostomata Odhner, 539.
Proteosoma, 656.
Protospirura Seurat, 573.
columbiana Cram, 575.
muricola Gedoelst, 538, 573, 584.
muris (Gmel.), 575.
Pseudoceocephalini, 390, 427, 432, 483, 435,
442.
Pseudocyphagogus Desbr., 397, 433, 437.
squamifer Desbr., 386, 397, 400, 437
Pseudorychodes Senna, 421, 435, 441.
prajclarus Kin., 389, 421, 422, 441.
Pseudothis pterocarpi Syd., 494.
Psidium guajava, 487, 511, 516, 527.
Psophocarpus, 518.
tetragonolobus, 518.
Psychotria luzoniensis, 495, 496.
Pterocarpus angalensis, 498.
indicus, 498. •
sp., 515.
782
Index
Puceinia citrata Syd., 480.
eongesta Berk. & Br., 480.
engleriana P. Henn., 480.
erebia Syd., 480.
heterospora Berk. & Curt., 480.
kuehnii (Krueg.) ButL, 480.
merrillii P. Henn., 480.
paullula Syd., 480.
philippinensis Syd., 480.
purpurea Cke., 480.
thwaitesii Berk., 480.
Pucciniaceae, 479.
Pucciniostele clarkiana (Barel.) Diet., 480.
Pueraria, 518.
sp., 532.
Pycnocrepis End., 294.
Pythiaceas, 518.
Pythium aoitumnale, 518.
debaryanum Hesse, 518.
esquiseti, 518.
vexans, 518.
Q
Quercus sp., 534.
Quinine, prophylactic, in inoculated avian
malaria, 347.
R
Raillietina Fuhrmann, 538, 548, 552.
celebensis (Janicki) Meggitt & Subra-
manian, 549, 585.
fluxa Meggitt & Subramanian, 586.
funebris Meggitt & Subramanian, 586.
garrisoni Tubangui, 538, 548, 549, 584-
586.
Ramularia catappae Rac, 530.
Rat, brown, worm parasites of, in the Phil-
ippine Islands, 537.
fleas, 556.
Rat-bite fever in the Philippines, 159.
Rhabdiasidas Railliet, 560.
Rhabdiasoidea Railliet, 560.
Rhabdospora synedrellae Sacc, 525.
Rhagadolobium bakerianum Sacc, 517.
Rhampholimnobia reticularis Alex., 23.
Rhipidia (Rhipidia) morionella Edw., 15.
Rhipidocarpon javanicum (Pat.) Theiss. &
Syd., 496.
Rhizopus artocarpi Rac, 520.
nigricans Ehrbg., 520.
Rhizosolenia, 92.
alata Brightw., 114.
alata Brightw. forma gracillima (Cleve)
Grun., 81, 114.
(alata var.) gracillima Clere, 114.
flaccida Castr., 114.
hyalina Ostenf., 115.
japonica Castr., 114.
robusta Norman, 115.
setigera Brightw., 81, 114.
sigma Schiitt, 115.
Rhododendron schadenbergii, 490.
sp., 490.
Rhopographella reyesiana Rehm, 500.
Rhopographus, 498.
Rhyticephalini, 433.
Rhytisma lagerstroemia Rabh., 517.
pongamiae, 517.
spurcarium, 495.
Ricinus communis, 518, 520, 522.
Rictularia Froelich, 538, 579, 580, 585.
tani, 585.
whartoni Tubangui, 538, 579, 584. 585.
Rictulariidse Railliet, 579.
Rictulariime Hall, 579.
RODA, ALFREDO PIO DE, see Oliver,
Leon and Roda.
RODRIGUEZ, JOSE, and FIDEL C. PLAN-
TILLA, The histamine test as an
aid in the diagnosis of early leprosy,
123.
Rosellinia bunodes (Berk. & Br.) Sacc, 502.
calami P. Henn., 502.
cocoes P. Henn., 502.
decipiens (Rehm) Theiss. & Syd., 502.
lamiprostoma Syd., 502.
megalosperma Syd., 502.
merrillii Syd., 502.
molleriana Henn., 502.
procera Syd., 503.
umbilicata Sacc, 503.
(Conimela) maquilingiana Rehm, 502.
(Tassiella) crustacea Rehm, 502.
(Tassiella) horrida Rehm, 502.
Rottboellia exaltata, 484, 486, 487, 499.
Rourea erecta, 491, 499.
Rubus moluccanus, 480.
RUSSELL, PAUL F., Avian malaria studies,
I. Prophylactic plasmochin in inocu-
lated avian malaria, 305 ; Avian ma-
laria studies, II. Prophylactic plas-
mochin versus prophylactic, quinine in
inoculated avian malaria, 347 ; Daytime
resting places of Anopheles mosquitoes
in the Philippines : First report, 639 ;
Avian malaria studies, III. The ex-
perimental epidemiology of avian ma-
laria ; introductory paper, 651.
Rhytisma, 495.
constellatum, 495.
S
Saccharum officinarum, 480, 484, 485, 496,
500, 519, 520, 528-530, 533.
spontaneum, 500, 525, 529, 535.
Sandoricum indicum, 519.
koetjape, 487, 519.
SANTOS, IRENE DE, AUGUSTUS P.
WEST, and P. D. ESGUERRA, Phil-
ippine pine-needle oil from Pinus in-
sularis (Endlicher), 1.
SANTOS, JOSJS K., Leaf and seed structure
of a Philippine Coriaria, 257.
Index
783
Sapindus saponaria, 505.
sp., 487.
Sarcinella raimundoi Sacc, 534.
Scamboneura O. S. 10, 271.
calianensis Alex., 10.
nigrotergata Alex., 9-11.
vittivertex Alex., 10.
Scarabseida?, 578, 759.
Scaunis striatus, 556.
Sceletonema costatum (Grev.) Cleve, 77.
Schizochora elmeri Syd., 500.
Schizceupsalis, 420.
Schizonema albicans V. Heurck, 113.
amplius V. Heurck, 113.
torquatum V. Heurck, 113.
Schizostachyum, 493, 497, 502, 503, 505-507,
510, 512.
acutiflorum, 500, 536.
lumampao, 510.
rotundifolium, 500.
sp., 500, 512, 533.
Schizothyrium aceris (P. Henn. & Lind.)
Pat., 515.
Schizotrachelus Lac, 429, 431, 435, 443.
agulaticeps Senna, 389, 429, 432, 443.
bakeri Kin., 389, 429, 432, 443.
bakeri Kin., f. concolor, 429, 443.
brevicaudatus Lac, 390, 430, 432, 443.
brunneus Kin., 390, 430, 432, 443.
consimilis Kin., 390, 430, 432, 443.
corpulentus Kin., 390, 430, 432, 443.
imbricellus Kin., 390, 430, 432, 443.
imitator Kin., 390, 430, 432, 443.
inconstans Kin., 390, 430, 432, 448.
Schlerospora spontanea Weston, 519.
Schneepia, 496.
hymenolepidis (P. Henn.) Theiss. & Syd.,
496.
SCH5BL, OTTO, An interpretation of the
laws of Brown and Pearce that govern
the course of treponematoses, 169 ;
Coexistent infection with yaws and
syphilis, 177 ; The prospects of vaccina-
tion and vaccine therapy in trepone-
matoses, 183.
Schroederella delicatula (Per.) Pavil., 99.
Stehroeteriaster cingens Syd., 481.
Scirpus grossus, 486.
Scirrhia, 494, 500.
bambusina Penz. & Sacc, 500.
luzonensis P. Henn., 500.
Scirrhodothis bambusina (Penz. & Sacc)
Theiss. & Syd., 500.
seriata Syd. & Butl., 500.
Sclerospora maydis (Rac) Butl., 519.
philippinensis Weston, 519.
sacchari Miyake, 519.
Sclerotinia nervisequia Schroet. v. bambusa-
cea Rehm, 516.
Sclerotium rolfsii Sacc, 536.
Scymnus sp., 221.
Septogloeum arachidis Rac, 529.
Septonema philippinum Sacc, 535.
Septoria palmarum Sacc, 525.
Septosporiella philippinensis Sacc, 525.
Serinus canarius, 656.
Sesamum indicum, 524, 526, 528, 529, 532.
orientale, 524, 526, 529, 532.
Sexes, determination of, in Leucopholis irro-
rata adults by use of antennae, 759.
Seynesia, 496.
aktoniae Rehm, 489.
calamicola, 496.
ipomoeae Syd., 489.
Shorea guiso, 195.
sp., 489.
Sida acuta, 487.
carpinifolia, 487.
javensis, 480.
Sideroxylon ferrugineum, 489.
sp., 489.
Skeleton of the timarau, 141.
SKVORTZOW, B. W., Pelagic diatoms of
Korean strait of the Sea of Japan,
95; Plankton diatoms from Vladivos-
tok Bay, 77.
Smilax bracteata, 480.
reticulata, 480.
Solanum melingena, 519, 522, 524, 532, 534.
tuberosum, 519.
Somatic segregation in double Hibiscus and
its inheritance, 627.
Sordaria oryzeti Sacc, 501.
Sordariaceae, 501.
Sorghum vulgare, 480, 484, 488, 500, 507,
534.
sp., 500.
Spegazzinia meliolae A. Zimm., 536.
ornata Sacc, 536.
Sphaeria, 492, 494, 496, 502, 506, 508.
Sphaeriaceaj, 501.
Sphaeriales, 501.
Sphaerioidacere, 520.
Sphaeriopsidales, 520.
Sphaerophragmium luzonicum Yates, 481.
Sphaerulina smilacincola Rehm, 505.
Spirillum minus, 165.
Spirochaeta morsus muris, 159, 165.
Spiroptera neoplastica Fibiger & Ditlevsen,
575.
Spiroxyinae Baylis & Lane, 573.
Spiruridae Oerley, 573.
Spiruroidea Railliet & Henry, 573.
Sponges, fresh-water, of the Philippine Is-
lands, 61.
Spongilla alba, 64.
dementis Annandale, 61, 65-69.
lacustris, 71.
microsclerifera Annandale, 61, 69.
philippinensis Annandale, 61, 62, 64-67,
69.
sceptrioides, 64, 68.
yunnanensis, 65.
Sporobolus elongatus, 534.
sp., 534.
Sporodesmium bakeri Syd., 5S4.
784
Index
Stagonospora varians Sacc, 525.
Stemonitis herbatica Peck., 85, 89.
splendens, 85.
splendens Host. var. flaccida Dister, 88.
Stenochlaena leptocarpa, 216.
smithii (Fee) Underwood, 216.
Stephanopyxis, 95.
appendiculata Ehrenb., 98.
campana Castr,, 98.
palmeriana (Grev.) Grun., 98.
palmeriana forma curta Alex., 98.
palmeriana var. japonica, 98.
turris (Grev. & Arn.) Ralfs, 98.
var. javanica Grun., 98.
Sterculia, 506.
Stereodermini, S90, 399, 431, 433, 434, 437.
Stereodermus Lac, 401, 434, 487.
fiavotibialis Kin., 387, 401, 437.
STEVENS, F. L., ed. Second supplement to
the list of the lower fungi of the
Philippine Islands, 479.
Stictidaceae, 517.
Stigmatodothis palawanensis Syd., 496.
Stigmelia manilensis Sacc, 535.
Stizolobium deeringianum, 481, 632.
niveum, 481, 532.
Streblus asper, 502, 511, 515.
Strobilocercus, 546.
fasciolaris Sambon, 546.
Strongyloidea Weinland, 567.
Strongyloides Grassi, 560.
papillosus (Wedl) Hall, 660.
ratti Sandground, 538, 560, 584.
stercoralis, 561.
Styiingomyia Loew, 294.
acuta Edw., 298.
armata Edw., 294, 297, 298.
ceylonica Edw., 295, 300.
claggi Alex., 294, 297.
colona Edw., 302.
ensifera Edw., 298.
flava Brun., 296.
flavocostalis Alex., 295, 299.
fumipennis Edw., 294, 296.
luteipennig Alex., 294-297.
mcgregori Alex., 294, 295.
montina Alex., 294, 296.
neocolona Alex., 295, 301.
nigrosternata Alex., 295, 299, 800.
tablasensis Alex., 295, 300.
taiwanensis Alex., 296.
SUMULONG, MANUEL D., The skeleton
of the timarau, 141.
Surirella gemma Ehrenb. var. ovata Skv., 116.
Symplocum whitfordii, 525.
Synchytriaceae, 517.
Synchytrium, 518.
aecidioides, 518.
decipiens, 518.
fulgens var. decipiens, 518.
Synedra, 77.
affinis Kutz. var. gracilis Grun., 81.
auriculata Karst., 112.
japonica Skv., 81.
Synedra — Conti nued.
koreana Skv., 112.
nitzschioides Grun., 110.
Synedrella nodiflora, 522, 525, 526.
Syphacia Seurat, 569.
obvelata (Rudolphi) Seurat, 538, 560,
584.
Syphaciinse Railliet, 569.
Syphilis, coexistent infection with yaws and,
177.
Kahn test in clinical, 225.
T
Tabernaemontana campanulata, 480.
polygama, 483.
Tachinida?, 129.
Taenia Linn., 546.
segyptiaca Bilharz, 556.
crassicollis Rudolphi, 546.
diminuta (Rudolphi), 553.
flavomaculata Leuckart, 553.
madagascariensis Leuckart, 585.
murina Dujardin, 556.
taeniaformis (Batsch) Wolffhugel, 538,
546, 584.
Taeniidse Ludwig, 546.
Taeniinae Stiles, 546.
Tsenioidea Zwicke, 546.
Talauma villariana, 506.
Tamarindus indica, 487.
Taphroderini, 433.
Teleneura Alex., 286, 287.
Telosma sp., 487.
Tenebrio molitor, 556.
Tenebrionidse, 578.
Tephrosticta ficina Syd., 509.
Terminalia catappa, 510, 530.
Tetracera sp., 495.
Tetrastigma, 517.
sp., 485.
Teucholabis (Teucholabis) confluenta Alex.,
36.
(Teucholabis) confluentoides Alex., 36.
(Teucholabis) majuscula Alex., 35, 36.
(Teucholabis) nigerrima Edw., 36.
Thalassiosira clevei Gran, 99.
gravida Cleve, 99.
hyalina (Grun.) Gran, 98.
nordenskioldii Cleve, 99.
Thalassiothrix antarctica Schimper, 80.
antarctica Schimper forma japonica
Skv., 110.
curvata Castr., 110.
frauenfeldi Cleve, 110.
frauenfeldi var. nitzschioides, 111.
frauenfeldii Grun., 80, 111.
frauenfeldii (Grun.) Castr., 111.
nitzschioides Grun., 80, 110.
nitzschioides var. javanica Grun., 111.
Thaumastoptera, 457.
(Taiwanita) calceata Mik, 457.
(Taiwanita) issikiana Alex., 467.
(Thaumastoptera) maculivena Alex., 456.
Index
785
Theobroma, 519.
cacao, 492, 508, 511, 519, 522, 530, 636.
Thunbergia grandiflora, 535.
Thyridaria calamincola Rehm, 5ll.
eminens Rehm, 511.
tarda Bancroft, 511.
Tilletiacese, 484.
Timarau, skeleton of the, 141.
Tipulidae from the Philippines (Diptera),
X, 9; XI, 269; XII, 447.
Tipulina?, 9, 269.
Torula dichroa Sacc, 535.
herbarum Lk., 535.
herbarum Lk. f. quatemella Sacc, 535.
Trabutia elmeri Theiss. & Syd., 501.
ficuum (Niessl.) Theiss & Syd., 501.
vernicosa Theiss. & Syd., 501.
Trachelizini, 390, 406, 431, 433, 434, 438.
Trachelizus Schoenh., 406, 484, 438.
bisulcatus F., 387, 406, 416, 438.
Trametes versatilis, 193.
Traversoa dothiorelloides Sacc. & Syd., 526.
excipuloides Sacc, 525.
excipuloides Sacc. & Syd. var. distans
Sacc. & Syd., 526.
Trema amboinensis, 489.
orientalis, 489.
sp., 489.
Trematoda Rudolphi, 539.
Trematodes, 538, 584.
Trematosphaeria maquilingiana Rehm, 503.
maquilingiana Rehm var. schizostachyi
Rehm, 503.
Trentepohlia, 473, 474.
brevifusa, 285.
riverai, 285.
(Anchimongoma) apoicola Alex., 475, 476.
(Anchimongoma) niveipes Edw., 476.
(Mongoma) sequialba Alex., 470, 471,
473, 474.
(Mongoma) aequinigra Alex., 471-473.
(Mongoma) distalis Alex., 284.
(Mongoma) luzonensis Edw., 471, 473,
474.
(Mongoma) majuscula Alex., 471, 473.
(Paramongoma) albitarsis (Dol.), 470.
(Paramongoma) banahaoensis Alex., 469,
470.
(Paramongoma) chionopoda Alex., 469,
470.
(Paramongoma) pusilla Edw., 470.
(Trentepohlia) festivipennis Edw., 475.
(Trentepohlia) laetipennis Alex., 474,
475.
(Trentepohlia) ornatipennis Brim., 474,
475.
(Trentepohlia) trentepohlii (Wied.), 27.
(Trentepohlia) venustipennis Edw., 475.
Treponema pallidum, 179, 241.
pertenue, 179, 241.
Treponematoses, experimental, comparative
serologic study of Vernes, Wasser-
mann, and Kahn reactions in, 241.
Treponematoses — Continued.
interpretation of the laws of Brown
and Pearce that govern the course
of, 169.
prospects of vaccination and vaccine
therapy in, 183.
Triblidiacese, 516.
Trichia asiatica Skv., 85, 90.
contorta, 85.
contorta Rost. var. inconspicua Lister,
90.
persimilis Karst., 85, 90.
varia Pers., 90.
Trichinella spiralis, 538.
Trichobelonium melioloides Rehm, 516.
Trichodolichopeza Alex., 270.
Trichonectria bambusicola Rehm, 493.
Trichophyton flava, 126.
Trichosanthes anguina, 523.
Trichosoma crassicauda Bellingham, 562.
muris decuman i Rayer, 562.
Trichosomoides Railliet, 562.
crassicauda (Bellingham) Railliet, 638,
562, 564, 584.
Trichosomoididse York & Maplestone, 562.
Trichosomoidinss Hall, 562.
Trichosphaeria bambusicola Rehm, 503.
Trichosporium coccidicola Sacc, 536.
Trichostrongylidse Leiper, 567.
Trichothyriacese, 490.
Trichothyrium orbiculare Syd., 489.
Trichuridae Railliet, 564.
Trichuroidea Railliet, 562.
Trionymus sacchari (CklL), 221.
Triumfetta sp., 485.
Tryblidiella mindanaensis P. Henn., 516.
rufula (Spreng.) Sacc, 516.
TUBANGUI, MARCOS A., Worm parasites
of the brown rat (Mus norvegicus)
in the Philippine Islands, with special
reference to those forms that may be
transmitted to human beings, 537.
Tuberculariaceae, 535.
Tychaeini, 433.
Tylophora fioribunda, 487.
perrottetii, 487.
U
Uleopeltis bambusina Syd., 496.
Ulocerini, 433.
Uncaria perrottetii, 487.
Uredinales, 479.
imperfecti, 482.
Uredo, 517.
arthraxonis-cillaris P. Henn., 483.
claoxyli Sacc, 483.
davaoensis Syd., 483.
desmium (Berk. & Br.) Petch., 483.
dioscoreae (Berk. & Br.) Petch., 483.
dioscoreae-alatae Rac, 483.
erythrinae P. Henn., 483.
fici Cast., 483.
786
Index
Uredo — Continued.
kuehnii (Krueg.) Wakk. & Went., 480.
manilensis Syd., 483.
mori, 482.
moricola, 482.
ochracea Diet., 483.
operculinae Syd., 483.
peckii, 517.
premnae Koord., 484.
vignae Bres., 484.
Uromyces, 518.
appendiculatus (Pers.) Lk., 481, 484.
deeringiae Syd., 481.
linearis Berk. & Br., 481.
mucunae Rabh., 481.
sojae Syd., 481.
vignicola, 418.
wedeliae P. Henn., 481.
Uatilaginacese, 484.
Ustilaginales, 484.
Ustilaginoidea ochracea P. Henn., 494.
virens (Cke.) Takafa., 494.
Ustilago andropogonia-aciculati Petch., 484.
fiagellata Syd., 484.
isaehnes Syd., 484.
manilensis Syd., 484.
eacchari Rabh., 484.
scitaminea (Rabh.) Syd., 484.
sorghi (Lk.) Pass., 484.
tonglinensis Tracy & Earle, 484.
Uvaria, 529.
rufa, 483.
Vaccination and vaccine therapy in trepo-
nematoses, 183.
Vallisneria, 71.
Valsaeeae, 510.
Valsaria citri Rehm, 511.
insitiva (de Not) Ces. & de Not, 511.
Vanilla sp., 503, 529.
VAZQUEZ-COLET, ANA, Rat-bite fever in
the Philippines, 159.
Vermicularia breviseta Sacc, 526.
capsici Syd., 526.
fallax Sacc, 526.
horridula Sacc, 526.
merrilliana Sac, 526.
sesamina Sacc, 526.
xanthosomatis Sacc, 526.
Vernes, Wassermann, and Kahn reactions,
comparative serologic study of, in ex-
perimental treponematoses, 241.
Vernonia vidali, 482.
Vigna sinensis, 518.
spp., 481, 484, 523.
Vladivostok Bay, plankton diatoms from,
77.
X
Xanthosoma sagittifolium, 526.
Xenopsylla* cheopis, 556.
Xylaria allantoidea Berk., 513.
castorea Berk., 514.
corniformis Fr„ 514.
euglossia Fr., 514.
grammica Mont., 514.
hypoxylon (L.) Grev. f. tropica Syd.,
514.
luzonensis Henn., 514.
nigripes (Klot.) Sacc, 514.
obvata Berk., 514.
plebeja Ces., 514.
tabacina (Kickx.) Berk., 514.
tuberosa (Pers.) Cke., 514.
Xylariacese, 512, 513.
W
WEST, AUGUSTUS P., see Cruz and West;
see also Santos, West, and Esgurrra.
Wood, decay of, in automobiles in the
Tropics, 189.
Worm parasites of the brown rat (Mus nor-
vegicus) in the Philippine Islands, 537.
Woroninella aecidioides (Peck.) Syd., 517.
dolichi (Cke.) Syd., 518.
psophocarpi Rac, 518.
puerariae (Henn.) Syd., 518.
Yaws and syphilis, 177.
YEAGER, CLARK H., Bored-hole latrine
equipment and construction, 681.
Ypselogonia Kin.. 422, 435, 441.
peregrina Kin., 389, 422, 424, 441.
Ypsilonia cuspidata Lev., 526.
Yucca aloifolia, 521.
Zea mays, 491, 501, 508, 519, 533, 534.
Zignoella (Trematostoma) nobilis Rehm, 503.
Zygomycetes, 520.
Zygosporium oscheoides Mont., 535.
O
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